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The CANADIAN 
FIELD-NATURALIST |. 


» 


Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada — 


Volume 119, Number 1 January—March 2005 


The Ottawa Field-Naturalists’ Club 


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Cover: Gray Jays, Perisoreus canadensis, feeding on White-tailed Deer, Odocoileus virginianus carcass near Ely, Minnesota. | 
See paper on weight-carrying ability and caching behavior by Lynn L. Rogers page 101-104. | 


THE CANADIAN 
FIELD-NATURALIST 


Volume 119 


2005 


Volume 121 
The Ottawa Field-Naturalists’ Club Transactions) 


Promoting the study and conservation 
of northern biodiversity since 1880 


THE OTTAWA FIELD-NATURALISTS’ CLUB 


OTTAWA CANADA 


The Canadian Field-Naturalist NI WARD 


Volume 119, Number 1 January—March 2005 


Chronology of Range Expansion of the Coyote, Camzs /atrans, 
_in New York 


HEATHER M. FENER!:2, JOSHUA R. GINSBERG?, ERIC W. SANDERSON?, and MATTHEW E. GOMPPER? 


'Center for Environmental Research and Conservation, Columbia University, New York, New York 10027 USA 
*Wildlife Conservation Society, 2300 Southern Boulevard, Bronx, New York 10460 USA 
3Department of Fisheries and Wildlife Sciences, University of Missouri, Columbia, Missouri 65211 USA 


Fener, Heather M., Joshua R. Ginsberg, Eric W. Sanderson, and Matthew E. Gompper. 2005. Chronology of range expansion 
of the Coyote, Canis /atrans, in New York. Canadian Field-Naturalist 119(1): 1-5. 


Coyotes (Canis /atrans) were historically restricted to central North America. In less than two centuries, however, Coyotes have 
colonized most of the continent, including much of northeastern North America. Better understanding causes and proximate 
mechanisms of this expansion requires a detailed understanding of how Coyotes colonized areas on a fine scale. We examined 
the establishment of Coyotes in the State of New York by collecting and analyzing reports of their first occurrence throughout 
the state over the past century, and creating a detailed map of range expansion. Coyotes first entered New York from the north, 
circled the Adirondack region prior to colonizing it, and then expanded southward and westward at ca. 78-90 km/decade. The 
revealed pattern lends little support to the hypotheses that the range expansion is attributable to translocations and releases, 
or that Coyotes were historically present in the region and only recently expanded in numbers. Rather, the data suggest a cor- 


relative relationship between anthropogenic land use and Coyote range expansion. 


Key Words: Coyote, Canis /atrans, range expansion, northeastern North America, landscape change, land use. 


The Coyote (Canis /atrans) is one of the most wide- 
ly disturbed and successful colonizing mammals in 
recent history. Traditionally restricted to the open grass- 
lands and praire ecosystems of Midwestern North 
America, today the Coyote is found throughout most 
of North America (Bekoff and Gese 2003). Eastern 
North America has been almost entirely colonized by 
Coyotes, which have become a numerically and ecolog- 
ically important member of the predator community 
(Parker 1995; Gompper 2002a, b). 

In northeastern North America, sporadic reports of 
Coyote-like canids in northern New England, New York, 
and the southeastern Canadian provinces of Ontario 
and Québec began in the early 1900s (Hilton 1978; 
Moore and Parker 1992; Parker 1995; Gompper 2002b). 
Through the 1930s and 1940s these reports became 
more frequent, and by the 1980s the Coyote was firmly 
established and widespread throughout the Northeast 
(Parker 1995). Several explanations may explain this 
broad and rapid range extension. Gray Wolf (C. Zzpus) 
extirpation throughout most of eastern North America 
in the 19" century is thought to have created an empty 
ecological niche ripe for exploitation (Parker 1995; 
Peterson 1996; Gompper 2002b). Extensive modifica- 
tion of the landscape through deforestation and agri- 
cultural development may also have facilitated range 
expansion (Lariviére and Créte 1992; Parker 1995). 


Importation and release of Coyotes into areas where 
they were previously nonexistent may also have aided 
their establishment in some regions (Parker 1995). 
Finally, some argue that Coyotes were always present 
in the Northeast, albeit in low numbers (Tullar 1992). 

Current maps depicting Coyote range expansion (e.g. 
Parker 1995) are ideal for visualizing continental-scale 
range expansion, but a finer scale resolution may assist 
in comparing the explanations for the current existence 
of Coyotes in the Northeast, and provide insights into 
the proximate mechanisms by which Coyote popula- 
tions expanded across the northeastern landscape. That 
is, how did these animals move through and establish 
themselves in the Northeast in such a relatively short 
period? Here we create a detailed (county-level) map 
of Coyote range expansion in the State of New York, 
and examine potential causes and mechanisms of this 
expansion. 


Methods 

To establish the Coyote colonization pattern and 
direction of their range extension in New York on a spa- 
tial and temporal scale, first occurrence reports and 
documented sightings of Coyotes were collected dating 
back to circa 1900. Because Coyotes were rare and 
undesirable newcomers in the region in the early 20" 
century, we believe these reports accuratly depict both 


2 THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Oe ee a 


ge 1940-1950 
1950-1955 
1955-1965 


fF!) 1965-1975 
he > ee OT O-2000 


BR 1 940-1945 (coyote-dog hybrid zone} 


FIGURE |. Coyote colonization pattern in New York between 1940 and 2000. A putative Coyote-feral dog hybrid population 
which existed along the northwestern edge of the Adirondacks is also indicated. 


presence and absence in particular areas and their move- 
ment across the state. County newspapers, regional out- 
door magazines (e.g., Adirondack Life, New York State 
Conservationist, and Fur, Fish & Game), and regional 
scientific journals (e.g. Mew York Fish and Game 
Journal) were examined for reports of Coyotes (time 
frame of search: 1880 — 2000), as were archives and 
collections of the American Museum of Natural History 
and the New York State Museum. 

County Clerks’ offices in northeastern New York 
(Albany, Clinton, Essex, Franklin, Fulton, Hamilton, 
Herkimer, Lewis, Jefferson, Madison, Montgomery, 
Otsego, Rensselaer, Oswego, Saratoga, Schenectady, 
Schoharie, St. Lawrence, Warren, and Washington coun- 
ties) were visited and canid bounty records and laws 
from the late 1800s and early 1900s were examined. 
In 2000 and 2001, field station, park, and preserve per- 
sonnel from throughout New York east of and includ- 
ing Oswego, Oneida, Madison, Chenango and Broome 
counties, where contacted to establish timing of Coyote 
establishment in their respective sites, and interviews 


(n = 38 individuals who were active with Coyote- 
related issues during the early stages of Coyote colo- 
nization) were conducted, with current and retired state 
biologists, wildlife technicians, university researchers, 
game wardens, professional trappers, hunters, taxi- 
dermists, fur buyers, and sheep farmers throughout 
eastern New York State (Fener 2001). 

From this data set a series of geographic information 
system data layers were created using ArcView/GIS 
mapping software (Environmental Systems Research 
Institute, Redlands, CA). Coyote reports were aggre- 
gated by county and by decade over the last 100 years, 
and then attributed to the county polygons in the GIS. 


Results and Discussion 

Coyotes became established in northern New York 
in the early 1940s (Figure 1). Prior to that there were 
occasional reports of Coyotes in the region; the earliest 
report is of a single individual from Franklin County 
in 1925 (Severinghaus 1974). In the early-mid 1930s 
Coyotes are again reported from Franklin County as 


2005 


FENER, GINSBERG, SANDERSON, AND GOMPPER: COYOTE RANGE IN NEW YORK 3 


FIGURE 2. Localities of reports of Coyotes that were likely introduced or released in New York. Symbols indicate decade in 
which Coyotes were reported. Circled symbols indicate known releases. Non-circled symbols were likely released 
animals based on proximity to known release sites. These original reports were all followed by extended periods of 
absence of Coyotes from the regions. Counties mentioned in text are numbered (1. Albany; 2. Bronx; 3. Broome; 4. 
Cayuga; 5. Chenango; 6. Clinton; 7. Essex; 8. Franklin; 9. Fulton; 10. Hamilton; 11. Herkimer; 12. Jefferson; 13. 
Lewis; 14. Madison; 15. Montgomery; 16. Oneida; 17. Oswego; 18. Otsego; 19. Rensselaer; 20. Saratoga; 21. Sch- 
enectady; 22. Schoharie; 23. St. Lawrence; 24. Warren; 25. Washington; 26. Westchester). Dark lines outline 
Adirondack State Park in northern New York and Catskill State Park in southern New York. 


well as neighboring Clinton County to the east. It is 
unclear, however, if these pre-1940s reports are valid 
(see below). Nonetheless, following the initial entry 
into New York from Québec or Ontario over the St. 
Lawrence River, Coyotes extended their range east 
into Vermont, and southwest along the St. Lawrence 
River towards Lake Ontario. This expansion occurred 
primarily along the periphery of Adirondack State Park, 
and in the early 1940s there was reportedly also a 
Coyote-domestic dog (“coydog”) hybrid zone along 
the northwestern periphery of the Adirondacks (Fener 
| 2001; see also Severinghaus 1974) (Figure 1). By the 
— early 1950s the range had expanded south along the 
New York/Vermont border and west back into New York 
State into the Albany area south of the Adirondacks. 
Coyotes did not show up in the Adirondacks in appre- 
ciable numbers until the late 1950s. In the 1960s, 
Coyotes were reported with increasing frequency in 


the Catskill region, and by the early 1970s had moved 
as far west as Cayuga County and as far south as West- 
chester County. In the 1980s Coyotes were commonly 
found throughout the state, excluding New York City 
and Long Island. In the 1990s Coyotes occurred in 
Bronx County of New York City, and a transient indi- 
vidual reached Central Park in Manhattan in 1999. 
As of 2002, only Long Island in southeast New York 
was not colonized by Coyotes. 

Coyotes colonized the ca 470 km North-South axis 
of New York in 60 yrs (1940-2000), or approximately 
78 km/decade. Excluding New York City, Coyotes col- 
onized the entire region (ca 450 km) by 1990 (90 kmm/yr). 
Periods of maximal range extensions include 1950- 
1960 when Coyotes expanded approximately 190 km 
southeast, and 1970-1980 when Coyotes expanded 
westwards by approximately 145 km. Estimating rates 
of range expansion in western New York are compli- 


4 THE CANADIAN FIELD-NATURALIST 


cated by the mixing of two expanding fronts. The 
original front of range expansion from northern New 
York via northern Ontario or Québec resulted in Coy- 
otes throughout eastern New York in the 1960s and west- 
central New York in the 1970s. A second colonization 
wave apparently entered New York from southern 
Ontario or northwestern Pennsylvania in the late 1960s- 
early 1970s (Figure 1). These two colonizing fronts met, 
such that by the 1980s all of western New York was 
occupied by Coyotes. 

Our results support the premise that Coyotes colo- 
nized New York through a range expansion from out- 
side the region, and not via expansion of a low-density 
population already existing in the state (Tullar 1992). 
Indeed, Parker’s (1995; pages 23-24) review of Coyotes 
in Ontario shows that the province was colonized in 
the 1920s and 1930s, suggesting Ontario, rather than 
Québec (colonized in the early 1940s; Parker 1995) 
as the source for the northern New York range expan- 
sion. There is minimal support for a colonization of 
New York prior to the 1930s, as most reports between 
1900 and 1930 were of released animals or occurred 
near sites of known releases (Figure 2; see also Fener 
2001). In addition, Coyotes remained absent from areas 
in which they were initiaily encountered in the 1920s 
and 1930s for several decades thereafter. The possible 
exceptions to this are reports from the mid 1920s-early 
1930s in far northern New York (see below). Two pri- 
mary areas of Coyote releases occurred. Four of five 
reports of Coyotes in west-central New York between 
1900 and 1930 can be linked to releases (Figure 2). In 
east-central New York there are six reports of Coyotes 
from the 1930s, four of which are directly linked to 
releases. Following these reports, however, Coyotes 
are not reported from these regions until the 1960s 
and the 1950s, respectively (Figure 1). Similar pat- 
terns have been observed in the southeastern United 
States, where some releases may have resulted in the 
establishment of isolated, local populations, but many 
other releases did not (Hill et al. 1987). 

Two additional releases in northern New York de- 
serve special attention because of their temporal and 
spatial proximity to the expanding Coyote front of the 
1940s. These releases occurred in Franklin (1934) and 
Jefferson (1941) counties. In Jefferson County, the 
release occurred after Coyotes had already colonized 
northern New York. Thus the release may have facili- 
tated the range expansion in counties east of Lake 
Ontario. The Franklin County release of 1934 may 
account for the observations of Coyotes in northern 
New York in the 1930s, as the species is not reported 
again from Franklin County until 1946. If this is the 
case — that is, that the 1930s observations of Coyotes 
in New York are based on introduced animals (see also 
Severinghaus 1974) — then the only report of a puta- 
tively naturally-colonizing Coyote in the state prior 
to the 1940s is a single animal from Franklin County 
in 1925 about 20 km south of the Québec border (Sev- 


Vol. 119 


eringhaus 1974). Southern Québec, however, was not 
colonized by Coyotes until 1944 (Parker 1995). It is 
therefore possible that this animal was also an escaped 
or released captive. 

The last documented Wolves in New York were 
killed in St. Lawrence and Franklin counties in the late 
1890s (Fener 2001), well before Coyotes entered the 
region. Thus the fine-scale pattern of Coyote range 
expansion offers little insight on the potential link 
between northeastern Wolf extirpation and Coyote 
range expansion. The absence of Wolves from New > 
York may, however, have increased the rate of range 
expansion in optimal Coyote habitat such as open 
areas, in turn allowing Coyotes to expand into subop- 
timal habitats such as heavily forested areas. Coyotes - 
generally prefer open or agricultural lands to heavily 
forested habitat (Post 1975; Samson and Créte 1997; 
Tremblay et al 1998; Créte et al. 2001). The pattern 
of range expansion in and around the Adirondacks of 
Northern New York suggests that Coyotes first ex- 
panded in agricultural areas followed by entry into 
more heavily forested regions 

Fener (2001) proposed that it was not the availability 
of open habitat per se that facilitated rapid expansion 
of Coyotes in New York, but rather the increase in 
abandoned farmlands which were in early succession- 
al stages of forest reestablishment when Coyotes en- 
tered the region. From 1920 to 1950, upwards of 2 mil- 
lion hectares of farmland were abandoned in New York. 
Farmland loss continues to the present, such that over 
the past century approximately 3 million hectares have 
been abandoned and left to regenerate naturally or been 
acquired and replanted by state reforestation programs 
(Alerich and Drake 1995; Stanton and Bills 1996). 
These habitats likely contain high densities of prey spe- 
cies for Coyotes. Thus the rapid expansion of Coyotes 
through New York may have been aided by entry into 
the region during a period when abandoned farmlands 
and early successional stages of forest were dominant 
landscape components. 


Acknowledgments 

This research was supported in part by funds from 
the Wildlife Conservation Society and the Center for 
Environmental Research and Conservation at Columbia 
University. We thank Justina Ray and Roland Kays 
for comments. 


Literature Cited 

Alerich, C. L., and D. A. Drake. 1995. Forest statistics for 
New York: 1980 and 1993. United States Department of 
Agriculture, Forest Service, Northeastern Forest Experi- 
ment Station. Resource Bulletin NE-132. 

Bekoff, M., and E. M. Gese. 2003. Coyote. Pages 467-481 
Wild Mammals of North America. Second edition. Edited 
by G. A. Feldhamer, B. C. Thompson, and J. A. Chapman. 
Johns Hopkins University Press, Baltimore. 

Créte, M. A., J.-P. Ouellet, J.-P. Tremblay, and R. 
Arsenault. 2001. Suitability of the forest landscape for 
coyotes in northeastern North America and its implica- 


2005 


tions for coexistence with other carnivores. Ecoscience 8: 
311-319. 

Fener, H. M. 2001. Coyote (Canis /atrans) colonization of 
New York State: the influence of human-induced landscape 
changes. M.A. thesis, Columbia University, New York. 
64 pages. 

Gompper, M. E. 2(002a. Top carnivores in the suburbs? Eco- 
logical and conservation issues raised by colonization of 
northeastern North America by coyotes. Bioscience 52: 
185-190. 

Gompper, M. E. 2002b. The ecology of Northeast coyotes: 
Current knowledge and priorities for future research. 
Wildlife Conservation Society Working Paper 17: 1-48. 

Hill, E. P., P. W. Sumner, and J. B. Wooding. 1987. Human 
influences on range expansion of coyotes in the southeast. 
Wildlife Society Bulletin 15: 521-524. 

Hilton, H. 1978. Systematics and ecology of the eastern coy- 
ote. Pages 210-228 7 Coyotes: biology, behavior, and man- 
agement. Edited by M. Bekoff. Academic Press, New York. 

Lariviére, S., and M. Créte. 1992. Causes et conséquences 
de la colonisation du Québec par le coyote. Québec Mini- 
stére du Loisir, de la Chasse et de la Péche, Direction géné- 
rale de la faune. SP-1935. 

Moore, G. C., and G. R. Parker. 1992. Colonization by the 
eastern coyote (Canis /atrans). Pages 23-27 in Ecology 
and management of the eastern coyote. Edited by A. H. 
Boer. Wildlife Research Unit, University of New Bruns- 
wick, Fredericton. 

Parker, G. R. 1995. The eastern coyote. Nimbus Publishing, 
Halifax. 254 pages. 


FENER, GINSBERG, SANDERSON, AND GOMPPER: COYOTE RANGE IN NEW YORK a 


Peterson, R. O. 1996. Wolves as intraspecific competitors 
in canid ecology. Pages 315-323 1 Wolves in a changing 
world. Edited by L. N. Carbyn, S. H. Fritts, and D. Seip. 
Canadian Circumpolar Institute, University of Alberta, 
Edmonton. 

Post, R. A. 1975. An ecological study of the northern Tug Hill 
coyotes. M.S. thesis, New York State College of Environ- 
mental Science and Forestry, Syracuse. 93 pages. 

Samson, C., and M. Créte. 1997. Summer food habits and 
population density of coyotes, Canis /atrans, in boreal 
forests of southeastern Quebec. Canadian Field-Naturalist 
L L023 227-233: 

Severinghaus, C. W. 1974. Notes on the history of wild canids 
in New York. New York Fish and Game Journal 21: 117- 
123; 

Stanton, B. F., and N. L. Bills. 1996. The return of agricul- 
tural lands to forest: changing land use in the twentieth 
century. Department of Agricultural, Resource, and Man- 
agerial Economics, College of Agriculture and Life Sci- 
ences, Cornell University, Ithaca. 

Tremblay, J.. M. Créte, and J. Huot. 1998. Summer forag- 
ing behavior of eastern coyotes in rural versus forest land- 
scape: a possible mechanism of source-sink dynamics. 
Ecoscience 5: 172-182. 

Tullar, B. Jr. 1992. The eastern coyote: always a New York 
native. New York State Conservationist January-February: 
34-39. 


Received 26 June 2003 
Accepted 18 January 2005 


Nesting Behavior, Ecology, Seasonal and Geographic Distribution of the 
Sand Wasp, S7ctiella emarginata (Hymenoptera: Sphecidae)* 


FRANK E. KURCZEWSKI and HUGH F. BOYLE 


Environmental and Forest Biology, State University of New York College of Environmental Science and Forestry, Syracuse, 
New York 13210-2778 USA 


Kurcezewski, Frank E., and Hugh F. Boyle. 2005. Nesting behavior, ecology, seasonal and geographic distribution of the 
Sand Wasp, S“ctella emarginata (Hymenoptera: Sphecidae). Canadian Field-Naturalist 119(1): 6-15. 


The nesting behavior and ecology of Swcte//la emarginata are documented for the first time based on field studies made 
mainly at Canadian Forces Base Borden, Simcoe County, Ontario. Type of soil, natural community, temporary closure, mound 
leveling, orientation flight, prey transport, nest structure and dimensions, and kind and number of prey per cell are defined. 


Museum and field collection records support a geographic bridge from northern Michigan to the Atlantic Coast and dispel . 


the previously held notion of a disjunct distribution for this species. A late June-July-early August flight season is inferred 
from observations and collections made in Ontario, New York and Michigan. The nesting behavior and ecology of 8 emar- 


ginata and several other Sw#cre//a species from the western United States, Mexico and Florida are compared. 


Key Words: Sand Wasp, S7ctie/la emarginata, Noctuidae, Hesperiidae, Ontario. 


The tribe Bembicini includes moderate to large- 
sized, stout-bodied sand wasps that are often conspic- 
uously marked with yellow or white (Parker 1917). 
All species of Bembicini nest in the ground, usually 
in sand or gravel. The subtribe Stictiellina of the tribe 
Bembicini includes five genera in the Nearctic Region: 
Stictiella, Glenostictia, Microstictia, Xerostictia and 
Steniolia (Bohart and Gillaspy 1985). Swcte/la con- 
tains 13 species that range collectively from Mexico 
into southern Canada. 

Species of S7ctie//a attempt to level the mound of 
soil that accumulates in front of an entrance from bur- 
row excavation. The species temporarily close the 
entrance with soil following burrow excavation and 
then make an orientation flight before going in search 
of prey. Species of Szcve//a hunt and stock under- 
ground cells with adult Lepidoptera (moths, skippers, 
butterflies). S#ce//a nests are one-, two- or many- 
celled depending on the species. The number of prey 
per cell is often inversely related to prey size. Most 
Stictiella species practice delayed mass provisioning; 
1.e., they lay an egg on the first prey placed in the cell 
before other prey are put inside (Evans 1966). 

Stictiella emarginata (Cresson) is the most widely 
occurring species in the genus. It ranges from Baja 
California through the United States into southern 
Canada (Bohart and Gillaspy 1985). This species was 
illustrated as having a disjunct geographic distribution 
with separate populations extending from the Great 
Plains to the Pacific Ocean and from the Appalachian 
Mountains to the Atlantic Ocean. Swctella emarginata 
is the only congener found in the northeastern United 
States and southeastern Canada. 


*Contribution number 2005-1 of the Roosevelt Wildlife Station 
of the State University of New York College of Environmental 
Science and Forestry, Syracuse, New York. 


Stictella emarginata \s virtually unknown ecolog- 
ically and behaviorally. The species is seldom seen in 
the field or collected east of the Rocky Mountains. 
Nothing is known of its nesting behavior except for a 
single prey record, an adult Zzroa [Noctuidae] from 
the western United States (Gillaspy et al. 1962). Brad- 
ley (1908) described a sleeping aggregation of 8° emar- 
ginata from California. Our paper documents the nest- 
ing behavior and ecology of 8 earginara for the first 
time based on observations made mainly at a single 
locality in southern Ontario. Our paper redefines the 
geographic and seasonal distribution of 8 earginata 
in the northeastern United States and southeastern 
Canada. 


Methods 

Nearly all field research on S. emarginafa was done 
at Canadian Forces Base Borden, Simcoe County, 
Ontario. We spent 15 days in the field, sometimes from 
0530 to 2000 hrs (EDT), at this site. We made field 
observations of wasps on 26-28 July 1996, 13-14 July 
1997, 5-7, 17-19 July 1998, and 29 June-2 July 1999. 
We found no adult S: evarginara nesting at Base Bor- 
den on 11-12 August 1995, 28-30 June 1996, and 27- 
30 June 1997. 

Field observations covering four hours were made 
on two females of 8. evzarginara at South Glens Falls, 
Saratoga County, New York, on 31 July 1993. We did 
not find any S! evarginatra at this locality during three 
visits on 18 July 1997, 14 July 1998, and 23 July 1999. 
We observed one female of this species for less than 
an hour at the Fort Drum Military Reservation, Jef- 
ferson County, New York on 22 July 1997. We did not 
find any §: emarginara at Fort Drum during 56 hrs of 
observation on 2-3 August and 18 October 1996 and 
3 April, 5-6, 12 July and 4 Oct 1997 (Kurczewski 
1998, 1999). The trips in October were made to ascer- 


6 


2005 


tain that there was no fall flight of 2" generation adults. 
The trip in April was made to ensure that the species did 
not winter and reappear in the spring in the adult stage. 

Two other sites with potentially favorable habitat for 
S. emarginata were investigated but the species was 
not found at either locality. Seven trips were made to 
the Rome Sand Plains, Oneida County, New York on 
27 June 1992, 28 July 1993, 13 July 1995, and 2 June, 
26 July and 10, 27 September 1997 (Kurczewski 1998). 
Time spent observing and collecting sphecid wasps 
at this site on Windsor loamy fine sand totaled 35 hrs. 
Four trips were made to the Albany Pine Bush, Albany 
County, New York on 29 June 1991, 18 July 1997, 14 
July 1998, and 23 July 1999. We spent 17 hours col- 
lecting and observing sphecid wasps at that locality 
on Colonie loamy fine sand. 

We spent over 1000 hours searching unsuccessfully 
for S. emarginata in sandy and gravelly habitats in the 
following states and counties in the northeastern Unit- 
ed States and southeastern Canada: INDIANA, Lake, 
LaPorte, Porter (21-22 June 1999); MASSACHUSETTS, 
Franklin (30 June 1991, 1 August 1993); MICHIGAN, 
Allegan, Alpena, Antrim, Arenac, Barry, Berrien, Cass, 
Cheboygan, Chippewa, Clare, Crawford, Emmett, Glad- 
win, Grand Traverse, Gratiot, losco, Kalamazoo, Kalka- 
ska, Lake, Luce, Manistee, Mason, Mecosta, Menomi- 
nee, Midland, Missaukee, Montcalm, Montmorency, 
Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Os- 
ceola, Oscoda, Otsego, Ottawa, Presque Isle, Roscom- 
mon, Van Buren, Washtenaw, Wexford (15-20 June 
1991; 9-12 June, 28 June-6 July, 7-10 August 1992; 
10-16 June, 8-12 July, 10-12 August 1993; 12-23, 26-27 
June 1994; 25-30 June 1995); NEw JERSEY, Atlantic, 
Burlington, Cape May, Ocean (19-21 August 1993, 
14-21 June 1995, 14-16 June 1996, 21-22 June 1997, 
24-25 June 1998); NEw York, Herkimer, Lewis, Onei- 
da, Onondaga, Oswego, St. Lawrence, Suffolk (3-5 
July 1994; 2-5 August 1996; 10 July 1997; 22, 25-27 
June 1998); Onto, Fulton, Lucas (28 June 1994; 14- 
20, 23-25 June 1999); ONTARIO, Elgin, Essex, Gren- 
ville, Grey, Haldimand-Norfolk Regional Municipality, 
Lambton, Northumberland, Prince Edward, Renfrew, 
Simcoe (20-27 June, 1-4, 12-25, 28 July 1996; 24-27 
June, 1-2 July 1997; 15-16, 19 July 1998); PENNSYL- 
VANIA, Erie (30 June 1994, 26 June 1999); QUEBEC, 
Kazabazua (31 July-1 August 1996), WISCONSIN, 
Adams, Bayfield, Douglas, Jackson, Juneau, Monroe 
(24-26 June 1994). 

Females of S! evarginara nested mainly on warm, 
sunny days. Wasps excavated burrows, searched for 
prey, provisioned cells, and closed nests at air tem- 
peratures of 20.6-28.9°C and sand surface tempera- 
tures of 28-51°C. At Base Borden, females worked at 
nests from 1005 to 2005 hrs. The last female to leave 
her nest in the evening to join a sleeping aggregation 
on vegetation departed at 2006 hrs on 5 July 1998. No 
nesting or any other wasp activity occurred after that 
time. We unearthed two nests at Base Borden that even- 
ing but did not find any adult wasps in them. Collection 


KURCZEWSKI AND BOYLE: NESTING, ECOLOGY, DISTRIBUTION OF THE SAND WASP / 


of this species at Fort Drum was made at 1515 hrs at 
an air temperature of 26.7°C. At South Glens Falls, 
wasps provisioned nests and excavated burrows from 
1321 to 1531 hrs at sand surface temperatures of 44- 
49°C. 

All wasps from 1998 and 1999 field studies at Base 
Borden were color-coded by applying different col- 
ored paint to the thorax, except for two females col- 
lected as voucher specimens. All nests were flagged 
with numbered wooden stakes. Type of wasp activity 
was noted, described and quantified. Emphasis was 
placed on burrow excavation, temporary closure, mound 
leveling, orientation flight, prey transport, final closure, 
and evening activity. Wasps entering nests with prey 
were timed between consecutive provisioning trips, 
from entry to exit, and during temporary closure, lev- 
eling and orientation flights. 

The burrow and cell(s) of each nest were excavated, 
examined, measured and drawn. Burrow length and 
design, number of cells per nest, cell depth, number of 
prey per cell, and position of prey in a cell and egg on 
a prey was recorded in the field. Prey, wasp cocoons, 
fly maggots and puparia were removed from the cells, 
placed in individual vials according to nest and cell 
number, put in an ice cooler, transported to a laboratory, 
and weighed [wet] on a Mettler balance. The aggre- 
gate prey weight of each cell was summed. The prey 
Lepidoptera were then pinned, code labeled, and, later, 
hand carried to Tim McCabe, New York State Insect 
Museum, for generic and specific determination. One 
of the two voucher specimens of 8! emarginata was 
sent to Howard Evans, Colorado State University, for 
species confirmation. Miltogrammini cleptoparasites 
were compared with specimens in the State Universi- 
ty of New York College of Environmental Science & 
Forestry collection determined by Margery Spofford. 
Wasp and prey specimens were deposited in the New 
York State Insect Museum, Albany. 

Ecological communities and habitats were defined 
using Varga and Schmelefske (1992) for Canadian 
Forces Base Borden, Ontario, Reschke (1990) for Karn- 
er and South Glens Falls, New York, and Kurczewski 
(1998) for the Fort Drum Military Reservation, New 
York. Soil type for Base Borden was identified using 
the Soil Map of Simcoe County [Base Borden Area] 
(Soil Research Institute of Canada 1959). Soil types 
for Fort Drum, Karner and South Glens Falls, New 
York were identified from soil samples sent to Ed Stein, 
United States Department of Agriculture, National 
Resources Conservation Service. 

To fill in gaps in the geographic distribution of S: 
emarginata we examined specimens and collection 
records from the University of Guelph, Royal Ontario 
Museum and Canadian National Collection for Ontario, 
Cornell University, American Museum of Natural His- 
tory and New York State Museum for New York State, 
and National Museum of Natural History, Smithsonian 
Institution, Pennsylvania Department of Agriculture, 
Carnegie Museum of Natural History, and The Penn- 


8 THE CANADIAN FIELD-NATURALIST 


sylvania State University for the northeastern United 
States. 

Weather information for Canadian Forces Base Bor- 
den, Ontario was unavailable from the base and nearby 
Angus weather stations as they were not operational. 
In order to simulate weather conditions at our research 
site (latitude 44°16'N, longitude 79°55'W, elevation 
221 m) for May-June 1996-1999, we obtained temper- 
atures from weather stations within 15 km of the base: 
Alliston Nelson (latitude 44°9'N, longitude 79°52'W, 
elevation 221 m), Egbert Care (latitude 44°13'N, lon- 
gitude 79°46'W, elevation 252 m), and Essa Hydro (lat- 
itude 44°21'N, longitude 79°49'W, elevation 216 m), 
Ontario. We averaged May-June 1996-1999 tempera- 
tures from these weather stations and used these aver- 
ages in a discussion of S. emarginata seasonal distri- 
bution. 


Results 
Geographic distribution 

We searched 10 insect museums in Ontario and the 
northeastern_United States and found the following 
unreported specimens of SS! emarginata. ONTARIO— 
Regional Municipality of York, Toronto, August 1918, 


ll 


Vol. 119 


1; Dufferin County, Primrose, June 1955, D. H. Pen- 
gelly, 3; Hastings County, Sydney Field Station near 
Foxboro, 8 July 1970, J. L. McAlpine, | female; NEw 
York: Albany County, Center [Karner], 28 July 1870, 
J. A. Lintner, 1 female. These collection records cou- 
pled with those from Pennsylvania (Parker 1929), 
northern Michigan (O’Brien 1989), Simcoe [Canadian 
Forces Base Borden] County, Ontario (Kurezewski 
2000), and Jefferson [Fort Drum Military Reservation] 
and Saratoga [South Glens Falls] Counties, New York, 
when plotted on a map, reveal a contiguous, transcon- 
tinental population of S: emarginara in the United States 
and southern Canada (Figure 1). 


Habitat and soils 
Canadian Forces Base Borden 

Two females were observed nesting at Base Bor- 
den in 1996 and 1997 in a sandy two-track running 
through a Red Pine [Aimus resinosa Aiton|—Scotch 
Pine [Pinus sylvestris L.|—graminoid savanna at the 
end of an airport runway (Figure 2). This two-track 
trail was the focal point of our 1998 and 1999 field 
studies. It was kept open by intermittent military vehi- 
cle use. In the early to mid-19" century droughty Tioga 
loamy sand (Soil Research Institute of Canada 1959), 


FIGURE |. Geographic distribution of S7ctie/la emarginava in the northeastern United States and southeastern Canada. Black 
circles represent collection localities as follows: MICHIGAN: Ontonagon County, Bruce Crossing; Marquette County, 
Huron Mountain Club (O’Brien 1989); ONTARIO: Dufferin County, Primrose; Simcoe County, Canadian Forces Base 
Borden; Regional Municipality of York, Toronto; Hastings County, Sydney Field Station near Foxboro; NEW YORK: 
Jefferson County, Fort Drum Military Reservation; Saratoga County, South Glens Falls; Albany County, Center 
| Karner]; PENNSYLVANIA: Cumberland County, Carlisle Junction (Parker 1929). Inset depicts geographic distribution 
of 8: emarginata as illustrated by Bohart and Gillaspy (1985). 


2005 


We Seal 


TY ok 


6 

me 
area 
1 des 
ae > 


s 


—< 


KURCZEWSKI AND BOYLE: NESTING, ECOLOGY, DISTRIBUTION OF THE SAND WASP 9 


FIGURE 2. Nesting site of S7cve/la emarginata at Canadian Forces Base Borden, Simcoe County, Ontario. 


continuous pine plains, and periodic fires character- 
ized the area. The coarse loamy sand coupled with 
ground and crown fires supported large Red Pine and 
White Pine [Pinus strobus L.| forests, savanna and 
barrens (Varga and Schmelefske 1992). Military activ- 
ity on the Camp Borden Sand Plain in the 20" century 
maintained the savanna, barrens and sandy openings 
(Kurczewski 2000). 


Karner 

Stictiella emarginata records from 1870 exist in the 
form of a female specimen in the New York State Insect 
Museum, Albany [see above] and a male recorded in 
A List of the Insects of New York with a List of the Spi- 
ders and certain other Allied Groups (Bradley 1928). 
The locality of these collections is Center, an old name 
for Karner, a 19" century railroad stop in the heart of 
the Pine Bush between Albany, Albany County and 
Schenectady, Schenectady County, New York. Pine 
barrens, pine plains, and shrub savanna or pitch pine 
[Pinus rigida Miller]-scrub oak [Quercus spp.] bar- 
rens (Reschke 1990) still persist in preserved areas on 
Colonie loamy fine sand. Fires occur regularly every 
eight to 11 years (Benton 1976). Pitch Pine-shrub oak 
was a dominant plant community of the 19" century 
Pine Bush (Milne 1985). We found no S. emarginara at 
Karner probably because of extensive habitat destruc- 
tion and expansion of alien vegetation due to fire sup- 
pression. 


South Glens Falls 

Two S. emarginata females nested in 1993 in 
recently bulldozed Windsor loamy sand of a con- 
struction equipment parking lot. The wasps nested 
near the road in a 5B slope that was too loose for 
heavy vehicles. This loamy sand contained many 
pebbles. Vegetation was similar to that at Karner, 1.e., 
pitch pine-scrub oak barrens (Reschke 1990) with a 
preponderance of Pitch Pine, White Pine, shrub oaks, 
and Sweet Fern [Comptonia peregrina (L.) J. M. 
Coulter]. 


Fort Drum Military Reservation 

One female was netted while flying with a noctuid 
moth held beneath her body in an area of Plainfield 
sand. The Fort Drum site contained open grassland and 
woodland of Pitch Pine, White Pine, Red Pine, and 
shrub oaks bordered by Sweet Fern (Kurczewski 1998). 
Original land surveys from the 1790s indicate that 
the area was once covered with White Pine, Pitch Pine, 
and oak forests with some sandy openings (Kurczewski 
1997, personal observation). The site is now kept open 
by constant military activity. Three species of psam- 
mophilous sphecid wasps that prefer vast expanses of 
barren sand, Ammophila harti (Fernald), Bembix pal- 
lidipicta Smith, and Philanthus albopilosus Cresson, 
were collected at this site in addition to S. emargina- 
fa (Kurczewski 1998). 


10 THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |. Average temperature (°C) at Alliston Nelson, Egbert Care and Essa Hydro, Ontario weather stations, May-June 
1996-1999 and first observed nesting date during those years at Canadian Forces Base Borden, Ontario. 


1996 1997 
Alliston 15.30 17.10 
Egbert 14.35 13.65 
Essa 14.60 13.80 
Average 14.75 14.85 
1* Nesting 26 July* 13 July 


*First observations by us, not first nesting by wasps at site. 


Flight season 

Stictiella emarginatfa has one emergence of adults 
per year in the Great Lakes Region according to col- 
lection dates on museum specimens and field obser- 
vations. The flight season of this species encompasses 
the last few days of June, July, and, rarely, early August. 
At Base Borden adult development, emergence and 
flight season was influenced by May-June temperature. 
Wasps-nested earlier (5-7 July 1998, 29 June-2 July 
1999) following warmer springs (May-June 1998, 1999 
average temperature, 17.40°, 17.17° C, respectively) 
and later (26-28 July 1996, 13-14 July 1997) after cool- 
er springs (May-June 1996, 1997 average temperature, 
14.75°, 14.85° C, respectively) (Table 1). Similar phe- 
nology in 1998 and 1999 |S! emarginara broods at Base 
Borden was probably related to similar May-June aver- 
age temperatures for those years. Unsuccessful attempts 
to find adults of this species at Base Borden following 
cooler springs (28-30 June 1996, 27-30 June 1997) 
supports the contention that emergence is tied to May- 
June temperature. 


Nest spacing and false starts 

Four females of ° emarginara nested at Base Borden 
in 1998 in a sandy two-track, 2.5 m wide, with a grassy 
median, 0.9 m wide. Each track was about 0.8 m wide 
(Figure 2). Swctella emarginafa was uncommon at this 
site compared to other sphecid species. More than 24 
Tachysphex similis Rohwer, eight 7 sarsatus (Say), 
four 7 pechumani Krombein, 10 Plenoculus davisi Fox, 
12-14 Bembix americana (Lepeletier), and 18-20 P/i/- 
anthus politus Say nests were scattered through the area 
of the two-track where SS: emarginara nested. 

Seven active nests of S: emarginata occupied the 
two tracks over three days. One section of the N track 
had one active nest and 10 false starts in an area 4.5 m 
long and 0.4 m wide. The distance between adjacent 
false starts ranged from 13 to 102 cm (Mean = 41.4 + 
26.37, N = 14). A section of the S track had three active 
nests and 26 false starts in an area 3.9 m long and 0.8 m 
wide. The three active nests were 25, 33 and 30 cm 
apart, respectively. The distance between adjacent false 
starts ranged from 2 to 90 cm (Mean = 27.5 + 17.58, 
N = 52). Three other. nests were located 4 m farther 
west on the S track. These nests, of different ages and 
possibly belonging to a single female, were only 15- 


1998 1999 
17.90 IETS 
16.90 16.80 
17.40 16.95 
17.40 17.17 
5 July 29 June 


18 cm apart. Four active nests at Base Borden in 1999 
were spaced 1.2-7.1 m [Mean = 3.5 + 2.52] apart. 
Two females nested about | m apart in 1993 at South 
Glens Falls. 


Nesting behavior 
Burrow excavation 

Four females at Base Borden excavated burrows 
between 1035 and 2005 hrs in 1998. The wasps made 
two to eight false starts before remaining in one place 
and completing a burrow. One female made seven false 
starts in an area 0.5 m? before finishing a burrow. Three 
other wasps moved as far as 0.5-3.0 m between con- 
secutive false starts. One female at South Glens Falls 
started digging a new burrow at 1531 hrs at a sand sur- 
face temperature of 44°C after nesting elsewhere from 
1321 to 1530 hrs at sand surface temperatures of 45- 
49°C, 

Females searching for a place to dig walked on the 
sand, tapped the surface with their antennae, and dug 
with the mandibles. One wasp turned completely on 
her side while using the mandibles. She produced an 
audible buzzing sound for >1 min. After the mandibles 
loosened the sand crust, she used the forelegs in unison 
to fling the loosened sand backward. Her body, espe- 
cially the abdomen, moved synchronously up and down 
to allow the sand to pass underneath. 

At Base Borden and South Glens Falls, where the 
soil contained loamy sand mixed with gravel, females 
constantly removed pebbles from their excavations. 
One wasp at Base Borden pulled pebbles backward 
with the mandibles nine times. She pulled them to the 
top of the sand mound in front of her entrance. The 
pebbles rolled down the sides of the mound and, later, 
became incorporated in the leveling process. The wasp 
then moved from the top of the mound straight into 
the burrow flinging sand backward with the forelegs. 
Females that dug nests in loamy sand without gravelly 
inclusions did not remove pebbles or make intermittent 
buzzing sounds while loosening the pebbles under- 
ground. 

After removing a pebble, most wasps made a hover- 
ing flight while facing the open entrance. The female 
that removed nine pebbles from her excavation made 
a hovering flight following eight of the nine sand re- 
movals. Most hovering flights were 2-4 sec in dura- 


2005 


tion but some lasted 5-10 sec. After removing sand or 
a pebble, some females paused and cleaned their anten- 
nae, eyes and mandibles with the forelegs. 

One wasp at Base Borden removed sand from her 
burrow 20 times in 50 min, or an average of once every 
2.5 min. A second female removed 62 sand loads in 
63.5 min, or about one load per minute. A third wasp 
dragged backward 49 loads of sand in 50.5 min. 
Other females spent 69 and 60 min to excavate a bur- 
row. At the beginning of an excavation, females were 
inside their burrows loosening sand with the mandibles 
for as long as 5-6 min before moving sand onto the 
surface. Toward the end of a dig, wasps often spent less 
than a minute for this behavior. After clearing loose 
sand and pebbles from the burrow, females made a 
temporary closure of the entrance. They appeared in 
the opening headfirst flinging sand backward with the 
forelegs in unison while tamping the fill with the end 
of the abdomen. Three wasps took 15, 18 and 45 sec 
to temporarily close their entrances, resulting in one- 
third to one-half of the sand mound being put back in 
the opening. 

After closing, females leveled the mound of soil in 
front of the opening. They walked across the mound 
in various directions flinging sand backward beneath 
the body with the forelegs until the mound was, more 
or less, flattened. Wasps nesting in mixed sand and grav- 
el then placed pebbles on the fill with the mandibles 
interspersed with making hovering [orientation?] 
flights. This sometimes involved moving debris out of 
the way. The hovering flights, 10-50 cm high, were 
15-50 sec in duration and made 10 sec to 2.5 min 
apart. During mound leveling as many as 10 hovering 
flights were interspersed after which a wasp rested for 
several seconds and then flew away. Mound leveling in 
five females took 5.5-11.0 min (Mean = 8.2 + 2.22). 


Provistoning 

Sixteen hours over three days were spent at Base 
Borden in 1998 and 1999 observing females bringing 
prey to nests. Four wasps provisioned nests from 1044 
to 1820 hrs at sand surface and air temperatures of 
28-36° and 22-25°C, respectively. They spent 9-77 
min [Mean = 28.6 + 22.70, N = 18] between consec- 
utive returns to a nest with prey. Females flew into the 
nesting area from a distance of >10 m. Such a wasp 
circled slowly in flight holding the prey ventral side 
upward with the middle legs while producing an 
audible buzzing sound. She then landed on the sand 
mound directly in front of an entrance or as far away 
as 50 cm before making a second flight to the mound. 
Two females at South Glens Falls brought prey to 
their nests at 1321 and 1442 hrs at sand surface tem- 
peratures of 44°C and 49°C. 

Retaining their grasp of the prey’s body with the 
middle legs, wasps raked open the temporary closure 
using the forelegs in unison. Removal of a closure usu- 
ally took 5-10 sec unless the area had been disturbed. 
Then, some females walked straight into the burrow 


KURCZEWSKI AND BOYLE: NESTING, ECOLOGY, DISTRIBUTION OF THE SAND WASP 1] 


holding the prey as described. However, most wasps 
released the prey ventral side upward just inside the 
burrow with its abdomen projecting from the entrance. 
Such females then turned around in the burrow, grasped 
the prey by its front end with the mandibles, and 
dragged it backward down the tunnel. Females spent 
about 2 min to remove the temporary closure, deposit 
the prey inside, oviposi7, and remake the closure for the 
first prey for the cell. Not having to oviposit, a female 
taking subsequent prey into a nest spent only 15-45 sec 
[Mean = 31.6 + 13.24, N = 18] inside and remade the 
closure in 6-9 sec. She then flew off immediately toward 
the hunting grounds making an audible buzzing sound. 
Some females rested on the sand before flying away. 
Other wasps made an orientation flight, rested on the 
sand, and flew away. 


Final closure 

After placing the full complement of prey in a cell, 
a wasp filled her burrow with sand and leveled the area 
of the entrance. One female at Base Borden appeared 
headfirst in her entrance 3 min after placing prey in 
the nest, walked onto the sand mound, and began rak- 
ing sand backward into the opening with the forelegs. 
She repeated this behavior eight times at intervals of 
15-97 sec [Mean = 65.0 + 30.24]. This wasp made 
brief hovering flights, facing the entrance, following 
her third and fourth trips onto the mound to get sand. 
Near the end of the closure, the female raked sand back- 
ward from the sides as well as the top of the mound. 
Always facing away from the opening, she pulled sev- 
eral pebbles into the burrow with the mandibles and 
packed in sand with the end of the abdomen. She spent 
nearly 11 min to fill the burrow, flew a short distance 
away, and rested on the sand. The wasp returned to 
the area 1.5 min later and finished raking sand and 
pulling pebbles onto the filled entrance. She then flew 
away, 13 min after taking her last prey into the nest. 


Nest structure and dimensions 

Twenty-six nest entrances at Base Borden in 1998 
and 1999 were mainly ovoid in shape and measured 
10 x 15 mm. Below ground, the burrows were circular 
and 8 mm in diameter. Twenty-two burrows entered the 
soil obliquely at angles of 45-60B with the surface 
and terminated in one cell (Figure 3). One nest each at 
Base Borden in 1998 and 1999 was two-celled with 
cells 3 and 4 cm apart (Figure 3). 

Base Borden burrows were significantly longer in 
(Oo Sethan in 1999 [7 = 3.75, dc. — 24°F <n], 
ranging from 72 to 138 mm [Mean = 100.9 + 24.35, 
N = 11] in 1998 and 62 to 91 mm [Mean = 71.4 + 
12.36, N = 15] in 1999. The three oldest 1998 nests 
had the longest burrows of 133, 138 and 135 mm. Cell 
depth in 1998 and 1999 at Base Borden was not sig- 
nificantly different, including the three oldest nests 
with the longest burrows [7= 0.51, df = 24, P = 0.62]. 
Cell depth in 1998 ranged from 43 to 73 mm [Mean 
= 53.3 + 11.90, N = 11] and in 1999 from 48 to 69 mm 


12 THE CANADIAN FIELD-NATURALIST 


FIGURE 3. One- and two-celled nests of S#ctiella emargina- 
fa as seen in side view. Sand mound is stippled. Scale 
refers to both nests. 


[Mean ='5 1:4 + 3.56, N = 15}; ‘Cell size [IN = 26] at 
Base Borden varied regardless of year: length [Range 
= 14-22 mm, Mean = 19.0 + 1.71], width [Range = 
13-15 mm, Mean = 14.4 + 0.75], height [Range = 12- 
15 mm, Mean = 13.1 + 0.94]. 


Cell contents 

From five to nine prey [Mean = 6.9 + 1.07, N = 18] 
were stored in fully provisioned cells at Base Borden. 
Individual prey weight ranged from 60.9 to 126.0 mg 
[Mean = 86.9 + 17.08, N = 132]. Two wasps from Base 
Borden [voucher specimens] weighed 147 and 151 mg. 
One noctuid prey from Fort Drum weighed 93 mg. Ag- 
gregate prey weight per cell from Base Borden ranged 


TABLE 2. Species of prey Lepidoptera of S#ctie/la emarginata. 


Family and species of prey 


NOCTUIDAE 
Apamea amputatrix Fitch 8 
Apamea ophiogramma (Esper) | 
Calophasta lunula (Hufnagel) pd 
Chytonix sensilis Grote 43 
I 
2 
Euxoa scandens (Riley) 3 
Euxoa incaltida (J.B. Smith) | 
Lacinipolia vicina (Grote) 14 
Nedra ramosula (Guenee) 21 
HESPERIIDAE 


Polites themistocles (Latreille) 8 


Number of specimens 


Vol. 119 


from 456.8 to 757.9 mg [Mean = 581.6 + 126.43, N 
=aSy. 

One cocoon from Base Borden, 69 mm deep, was 
7 mm wide and 17 mm long, weighed 358.3 mg, and 
reared a male wasp. Two cocoons from Base Borden, 
72 and 73 mm deep, were each 8 mm wide and 19 mm 
long, weighed 567.5 and 604.6 mg, respectively, and 
reared female wasps. 

Of 132 prey in the cells, 78 (59.1%) were positioned 
head inward and ventral side upward. Fifty-four (43.5%) 
of the prey were placed head inward and on the side. 

Females preyed on adults of seven species of Noc- 
tuidae and one species of Hesperiidae. Of 132 prey 
specimens, 124 (93.9%) belonged to the family Noc- 
tuidae and eight (6.1%) belonged to the family Hes- 
periidae (Table 2). Some prey species were associated 
with barren sandy and/or fire- adapted habitats. Prey 
individuals were probably captured on flowers or low 
growing herbaceous vegetation. 


Leg 

Stctiella emarginata eggs from four females at Base 
Borden in 1998 were approximately 4 mm long and 
1 mm wide [N = 11]. Two recently laid eggs weighed 
2.0 mg while a one day-old wasp larva weighed 2.15 mg. 
Eggs or 1* instar larvae were attached to the base of the 
prey’s abdomen near its connection to the thorax or to 
the side of the thorax. The egg extended longitudinally 
along the side of the thorax toward the head. Eggs or 
1‘ instar larvae were affixed about equally to either 
the left [N = 14] or right [N = 12] side of the prey. 

A wasp’s egg was affixed to the initial prey in the cell 
in eight nests at Base Borden. The position of the prey 
on which the wasp’s egg or larva was affixed changed 
as additional prey items were added to the cell. In one 
incomplete cell with three moths, an egg was attached 
to the prey that was farthest in. In one cell with seven 
moths, a small larva was feeding in place on the fourth 
prey from the back end. In other cells, older larvae had 
moved around and were feeding on prey near the front 
end as these were the only prey left with nutrient value. 


Source 


C.F.B. Borden 
C.FB. Borden 
C.F.B. Borden 
C.F.B. Borden 

Ft. Drum 

South Glens Falls 
C.F.B. Borden 
Gillaspy et al. 1962 
C.F.B. Borden 
C.F.B. Borden 


C.F.B. Borden 


2005 


Mortality 

Five of 26 (19.2%) cells at Base Borden were afflict- 
ed with cleptoparasitic miltogrammine maggots or pu- 
paria. This included a one-celled nest in 1998 and two 
two-celled nests in 1998 and 1999. The single-celled 
nest held five paralyzed noctuids, no wasp’s egg, two 
large maggots and two small maggots. The different 
sizes of the maggots indicate that the flies attacked at 
two different stages in the provisioning of the cell, per- 
haps a day apart. Both cells in the 1998 two-celled nest 
each held two miltogrammine puparia and six pairs of 
noctuid wings. The puparia in one cell were 5.5 mm 
long and 2 mm wide and weighed 21.9 and 21.3 mg, 
respectively. One cell of a two-celled 1999 nest con- 
tained seven pairs of noctuid wings and four milto- 
grammine puparia. The other cell held seven paralyzed 
noctuids, a small wasp larva weighing 2.15 mg, and 
four miltogrammine fly maggots. No maggots were 
reared to adult flies. Two puparia reared a male and fe- 
male Sphixapata vigilans Allen |Sarcophagidae: Mil- 
togramminae]. This fly species was observed on sand 
near S. emarginata nests and pursued provisioning 
wasps as they landed and entered nests. 


Discussion 

The geographic distribution of 8° emarginata, as 
illustrated by Bohart and Gillaspy (1985), is disjunct 
(Figure 1). Specimens collected in the 1980s from the 
Upper Peninsula of Michigan brought these populations 
closer together (O’Brien 1989) and led us to believe 
that the disjunct distribution of 8. emarginata was an 
artifact resulting from lack of collection records. Col- 
lection records from southern Ontario and Upstate New 
York field studies and museum specimens indeed bridge 
the gap between northern Michigan and the Middle 
Atlantic Region and provide evidence for a continuous, 
transcontinental geographic distribution for S. ear- 
ginata. 

StictieHa emarginata has a flight period from early to 
mid-summer in southeastern Canada and northeastern 
United States. Collection dates for adults of this species 
from eight localities in northern Michigan, southern 


TABLE 3. Nesting characteristics of species of S7ctie//a. 


KURCZEWSKI AND BOYLE: NESTING, ECOLOGY, DISTRIBUTION OF THE SAND WASP 13 


Ontario and Upstate New York range from 29 June 
[1999] to 31 July [1993]. There is one record from Tor- 
onto, Ontario labelled simply “August.” Adult develop- 
ment, emergence and nesting at Canadian Forces Base 
Borden, Ontario seem to be tied to May-June temper- 
ature. Wasps nested earlier (5-7 July 1998, 29 June-2 
July 1999) with warmer May-June 1998, 1999 average 
temperatures (17.40°, 17.17°C) and later (26-28 July 
1996, 13-14 July 1997) with cooler May-June 1996, 
1997 average temperatures (14.75°, 14.85°C). 

Nesting information is available for six S”#cre//la 
taxa: ca//ista Parker, emarginata, evansi Gillaspy, /or- 
mosa (Cresson), p. pulchella (Cresson), and pulchella 
serrata (Handlirsch) (Table 3). Females of all species 
make a temporary closure of the nest. All S7ctie//la 
attempt to level the mound of soil in front of their 
entrance before flying off in search of prey. There is 
contradictory information about the leveling behavior 
of S: formosa. Lin in Gillaspy et al. (1962) observed 
a female “throwing...soil” behind her as she walked 
across the soil mound and entered the burrow. This 
wasp also walked across the mound in a different 
direction throwing soil before turning and reentering 
her entrance, yet Lin stated that the mound was “never 
leveled.” 

A clearly audible, high-pitched buzz accompanies 
an orientation flight in Swc/e//a species (Alcock and 
Gamboa 1975; Evans 1966; Gillaspy et al. 1962; 
Krombein 1964; this study). In S! pul/chella serrata and 
S. formosa females followed potential prey as they 
visited low herbs, grasses, and flowers and then cap- 
tured an individual with one quick strike before car- 
rying it away in flight (Gillaspy et al. 1962). Swctella 
pulchella serrata, S. p. pulchella and S. evansi capture 
moths, 8: ca//ista moths, skippers and butterflies, S- 


Jormosa skippers and butterflies, and S. emarginata 


moths and skippers (Alcock and Gamboa 1975; Evans 
1966; Gillaspy et al. 1962; Krombein 1964; Parker 
1917; this study)(Table 3). One female of 8! cosniculata 
Mickel is pinned with a noctuid moth (Bohart and 
Gillaspy 1985). Some cells contained only one prey 
species or genus [formosa, p. pulchella\. In other cells 


Species of S7ctiella* 


Characteristics callista emarginata 
Temporary closure + + 
Leveling behavior + + 
Burrow length (cm) 68 6-14 
Cell depth (cm) 40 4-7 
Number of cells/nest l 1-2 
Number of prey/cell > 5-9 
Prey type LIN? HN 


eVanst Jormosa pulchella serrata 
+ + + 
+? + + 
15-24 20 25-32 15-28 
11-16 8-14 12-20 10-14 
| 5-17 1-2 l 
15 7-11 19 12->21 
SG HML SN CEOPSTY 


*pulchella and serrata are subspecies of Stictiella pulchella. 
**Prey families of Lepidoptera are abbreviated as follows: C, Crambidae; E, Epipaschiidae; G, Gelechiidae; H, Hesperiidae; 
L, Lycaenidae; M, Nymphalidae; N, Noctuidae; O, Olethreutidae; P, Pyralidae; S, Pyraustidae; T, Tortricidae; Y, Phycitidae. 


14 THE CANADIAN FIELD-NATURALIST 


the species, genera and families were mixed within 
the known range of prey types [/ovmosa, p. pulchella, 
pulchella serrata, emarginata, evansi\. 

Prey of S. emarginata vary in their feeding habits and 
are probably captured in different places. Ca/ophasia 
/unula (Hufnagel) feeds on Butter-and-eggs [Zimaria 
vulgaris Miller]. Apamea ophiogramma (Esper) feeds 
primarily on wetland grasses. Apamea amputatrix Fitch 
and Polites themistocles (Latreille) are also grass-feed- 
ing but may lay their eggs on non-grasses. Adults of 
P. themistocles are commonly seen on flowers along 
country roads. Nedra ramosula (Guenee) is host-spe- 
cific on St. John’s-wort [Hypericum perforatum L.}. 
Lacinipolia vicina (Grote) is a general feeder on low- 
growing herbaceous plants. Chyfonix sensilis (Grote) 
feeds on a variety of fungi 15-20 years after a fire. 
Euxoa scandens (Riley) occurs in areas of dry, sandy 
soil and feeds on dicotyledons (Layberry et al. 1998; 
Rings and Downer 2001; T. L. McCabe 1999 person- 
al communication). 

Prey transport in Szctie//a species is by flying, the 
wasp holding the prey head forward and ventral side 
upward with the middle or middle and hind legs. Size 
of prey may influence whether the hind legs assist the 
middle legs in prey carriage (Krombein 1964). Prey 
Hesperiidae of 8. fovmosa were carried with the wings 
partly spread while prey Pyralidae of S: pulchella 
serrafa Were transported with the wings folded tight- 
ly against the body (Gillaspy et al. 1962; Krombein 
1964). Prey Noctuidae and Hesperiidae of S. emar- 
ginata were carried both ways (this study). The prey 
is held only with the middle legs when removing the 
temporary closure and entering the burrow. 

Final nest closure has been described only in S:- 
emarginata (this study). Such behavior involves the 
wasp walking onto the sand mound and flinging soil 
backward into the open burrow. She rakes backward 
the loose soil using the forelegs in unison as she backs 
into the burrow and occasionally tamps this soil with 
the end of the abdomen. She levels the area of the en- 
trance after filling the burrow flush. The entire process 
takes only one-fourth as long as burrow excavation. 

There is much variation among S/c/e//a species in 
nest dimensions (Alcock and Gamboa 1975; Evans 
1966; Gillaspy et al. 1962; Krombein 1964; this study) 
(Table 3). Although excavated in sandy soil, burrows 
and cells of 8 emarginafa were much shorter and 
shallower than in other species. The number of prey 
per fully provisioned cell in Swcve//a species is highly 
variable (Table 3). Swctella emarginata is unique in 
that only five to nine prey are stored per fully provi- 
sioned ceil. Wasp cocoons recovered from old 8 emar- 
ginata cells were enswathed with only six or seven pairs 
of noctuid wings (this study). 

Although S/cte//la species practice mass or delayed 
mass provisioning, there is one record for S: pulchella 
serrata that is suggestive of progressive provisioning. 
In that observation, a one-third grown larva was found 
in an incompletely provisioned cell with 21 moths. 


Vol. 119 


The nest had not received a final closure (Krombein 
1964). There are records of delayed mass provision- 
ing in S: callista and S. evans in which larval wasps 
occupied incompletely provisioned cells (Alcock and 
Gamboa 1975; Evans 1966). These cells and that of 
S. pulchella serrata were provisioned for parts of at 
least two days. 


Acknowledgments 


Howard Evans (now deceased), Colorado State Uni- - 


versity, and an anonymous reviewer critically read the 
manuscript. Tim McCabe, New York State Museum, 
identified the prey Lepidoptera, furnished host plant 
information, and located a historical specimen of S- 
emarginata. Matthias Buck and Steve Marshall, Uni- 
versity of Guelph, and Lubomir Masner and Jeff Skev- 
ington, Canadian National Collection, furnished south- 
ern Ontario collection records. Gary Hevel, National 
Museum of Natural History, Smithsonian Institution, 
Washington, gave us collection information from labels 
on specimens of S! emarginata from Pennsylvania. 
Bill Huff, Canadian Forces Base Borden, provided 
access to and lodging at the base and apprised us of 
weather conditions and potential wasp emergence. 


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KURCZEWSKI AND BOYLE: NESTING, ECOLOGY, DISTRIBUTION OF THE SAND WASP Bs 


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Received 19 June 2002 
Accepted 4 March 2005 


Environment and Distribution of Age O Fishes in River Canard, | 
a Lowland Ontario River 


JOHN K. LESLIE and CHARLES A. TIMMINS 


Department of Fisheries and Oceans, Great Lakes Laboratory for Fisheries and Aquatic Sciences, 867 Lakeshore Road, 
Burlington L7R 4A6 Canada 


Leslie, John K., and Charles A. Timmins. 2005 Environment and distribution of age 0 fishes in River Canard, a lowland Ontario 
river. Canadian Field-Naturalist 119(1): 16-25. 


Age 0 fishes were collected to determine occurrence, relative abundance and species composition at three sites in River 
Canard, Ontario in spring-autumn, 1994-1995. This small lowland river (mean annual discharge, 3.2 m* s"!) has variable 
flow during fish spawning and early nursery periods, high suspended particulate load, and sparse rooted vegetation and 


other physical cover. Forty-two taxa (12 families; 24 544 specimens) collected with beach seines and a plankton net repre- - 


sented a wide range of reproductive strategies and a diverse taxocene. Gizzard Shad Dorosoma cepedianum (67% of total 
catch), Orangespotted Sunfish Lepomis humilis (8%), and Brook Silverside Labidesthes sicculus (6%) were the most abun- 
dant species. Environmental conditions were such that fish attained autumnal lengths comparable to species in various sys- 


tems throughout the ecoregion. 


Key Words: age 0 fishes, abundance, growth, turbidity, lowland river, River Canard, Ontario. 


The largest number of freshwater fish species in 
Canada inhabit tributary streams, marshes and near 
shore waters of Lake St. Clair and western Lake Erie. 
Despite more than 200 years of human interference 
throughout the area, the fish community maintains a 
surprisingly wide spectrum of feeding and reproduc- 
tive guilds. Many species considered of minor impor- 
tance to local and regional economies utilise an exten- 
sive network of low gradient, silt-laden streams and 
drainage systems that lace the landscape (Leslie and 
Timmins 1990; 1998a, 1998b; Leslie et al. 1999). The 
eventual movement of fishes from these waters to larger 
systems such as the Detroit River, Lake St. Clair, and 
Lake Erie contributes important fauna and nutrients 
to the lower Great Lakes basin (Herdendorf 1987). 

Because many aspects of fish reproduction and early 
life history have large knowledge gaps, success of 
fish habitat conservation, enhancement and develop- 
ment rests largely on synopses of general ecology and 
anecdotal information. Nevertheless, in Canada, the 
Fisheries Act (Canada Department of Fisheries and 
Oceans [DFO] 1984) requires protection and enhance- 
ment of spawning, nursery, and residence habitat of 
all species, regardless of their economic importance 
and despite limited commitment to their study. More- 
over, a thorough knowledge of the ecosystem is nec- 
essary for proper fisheries management and to improve 
our ability to determine the capacity of the system 
for sustainable development (Haffner 1992). 

This paper describes the relative contribution and 
seasonality of age O fishes at three locations in River 
Canard, a tributary to the Detroit River. This study 
intended to provide information on features of early 
life history of species in small tributaries, and is the 


first such investigation in a lowland river in the lower 
Great Lakes. 


Study Area 

River Canard (42°10'30"N; 83°05'00"W) is a base 
gradient (<0.4 m km’), meandering stream draining 
approximately 159 km? in the St. Clair flats, Ontario. 
It is about 25 km long, with a mean annual discharge 
of 3.2 m’ s! (Ontario Ministry of Environment and 
Energy [MOEE] 1994). The drainage area is predom- 
inantly heavy textured, with poorly drained soils of 
dark grey spleisolic, with Devonian limestone dominat- 
ing the bedrock (Chapman and Putnam 1984). Water 
quality is strongly influenced by runoff from intensely 
cultivated cropland that has a legacy of pesticide, her- 
bicide and fertiliser application. In addition, the eco- 
system is subjected to detrimental effects of intense 
vehicular traffic on several major highways. Agricul- 
tural and municipal drainage ditches connect River 
Canard with Lake Erie through Big and Cedar creeks 
and Lake St. Clair through the Pike Creek watershed 
(Figure 1). 

Winner and Hartt (1969) described aspects of the 
aquatic biota of the river, exclusive of fishes. The 
rotifer Brachinus sp., an indicator of eutrophic systems, 
dominated the zooplankton community. In the lower 
river, common submersed vegetation included pond- 
weed Foramogeton sp., Coontail Ceratophyllum demer- 
sum, Eurasian milfoil A74yriophyllum sp., stonewort 
Chara sp., and Eelgrass Va//isneria americana. Dom- 
inant emergent plants were cattail 7ipha latifolia, T. 
angustifolia, bur-reed Sparganium sp., bulrush Scirpus 
sp., and sedges Carver spp. (Winner and Hartt 1969). 
This community of plants prevails, with the addition 


2005 


of American Lotus WVe/umbo lutea, an exotic emer- 
gent established near the outlet to the Detroit River. 

The general character of the river does not change 
drastically along its course; 1.e., there are no pools or 
riffles, or vegetated backwaters. However, three sam- 
pling sites were chosen based on slight differences in 
physical and biological characteristics. Two sites (mid- 
dle and upper reach) on the main course of the river 
and one (lower reach) near the confluence of a tribu- 
tary to River Canard (Figure 1) were sampled for 
fishes. The upper reach (U: 42°07'36"N; 82°58'26"W), 
located about 14 km from the effluent, flows through 
pasture and scrubland. Stream width, typically 3 m, 
and depth, 0.2-0.5 m, vary according to seasonal rain- 
fall and flood control measures. In drought conditions, 
the upper reach is dry. Gravel substrate predominates 
in the centre, with sand and clay near gently sloping 
banks. Emergent plants consist mainly of Broad-leaved 
Cattail 7ipha latifolia, Common Reed Phragmites 
australis, and Great Bulrush Scivpus validus. Sparse 
riparian vegetation consists of small shrubs; e.g., 
Hazelnut Corylus americana and Dogwood Cornus 
racemosa. Partially submersed terrestrial grasses at 
the stream margin provide minimal cover and associ- 
ated food for fishes. Crayfish Ovconectes propinquus 
are abundant at all times. At least 20 drainage ditches 
are connected to the river between its origin and the 
upper reach sampling site. 

The middle site (M: 42°10'72"N; 83°02'05"W) is 
situated on a tributary that meanders through crop- 
land and pasture 0.2 km east of its confluence with 
River Canard. Midstream depth and width were | m 
and 11 m, respectively, and the substrate consists of 
fine gravel and sand in the centre, with steep clay banks. 
No rooted aquatic plants are established, whereas trees, 
such as hawthorn Cra/aegus sp., Red Maple Acer 
rubrum, and Cottonwood Populus deltoides, form a 
closed canopy from mid-spring to late autumn. 

In the lower reach (L: 42°10'29"N; 83°05'91"W), 
sampling took place 0.2 km from the Detroit River. 
At this location, River Canard is 230 m wide and about 
1.0 m deep. Fish collections were taken at the margin 
of American Lotus, where grey clay is overlain with 
alluvium. Submersed macrophytes are sparse and scat- 
tered among debris of human origin. Broad-leaved 
Cattail, Giant Bulrush, and Common Reed are abun- 
dant at the shore. 


Methods 

In 1994, sampling took place only in the lower 
reach. Routine fish collections were taken weekly or 
twice monthly from late June to November. In 1995, 
collections began in mid-April and continued weekly 
or twice monthly until October, then once in Novem- 
ber. Beach seines at the shore and a conical plankton 
net towed in open water were used to collect age 0 
fishes. Most collections were made with a bagged lar- 
val fish seine (length 4 m, width | m, 0.3-mm Nitex 
mesh), hauled 10-15 m parallel to shore at maximum 


LESLIE AND TIMMINS: AGE O FISHES IN RIVER CANARD I? 


| 
83° 06 W 


Michigan 


setti Lake St. Clair 
ras) 


Pike Ck _ 


N 
Ontario 
eole 23) 
5 km 
Canard River 
— 42° 06'N U 


Cedar Ck 
Lake Erie 


FiGuRE |. Map of study area, showing fish collection sites: 
L (lower reach), M (middle site), and U (upper- 
reach) in River Canard, Ontario in 1994-1995. 


wading depth of about 1.0 m. The employment of 
this sampler is unique. At the end of each horizontal 
sweep, the seine 1s rapidly arced vertically to just above 
the surface, where contents are washed into its centre 
(Leslie et al. 1983). Each collection was immediately 
replicated at least once in the same location on each 
date. Specimens were immediately fixed with 5-20% 
formalin. Additionally, a 6-m long, 1-m wide beach 
seine (6 mm mesh), used to provide ancillary infor- 
mation (species occurrence, seasonality) on age I* 
(juvenile and adult) fishes, swept a horizontal dis- 
tance of about 15 m on 2-6 successive sampling efforts. 
Twenty plankton tows were made in the lower reach 
to determine if species compositions in open water 
differed with those at the shore during usual peak 
occurrence of larvae in mid to late-June. Alternate 
tows were made upstream and downstream during 
the day over a constant distance of 230 m at speeds 
of 0.7-1.1 m/s (average, 0.9 m/s). A 0.4-m diameter, 
1.5-m long conical net (0.4 mm Nitex mesh) collect- 
ed fishes near the windward shore, the only habitat 
where depth was sufficient to perform linear hauls. 
The net was towed just below surface 10 m from the 
stern of an inflatable raft pushed by a small motor. 
Fish densities were expressed as number/100 m°. 
Sampling took place above a substrate of gravel, sand, 
and alluvium where maximum depth was 1.0-1.3 m. 
Jaccard’s coefficient of community Cc (Oosting 
1956) was used to compare longitudinal occurrences 
of fishes in the river. Cc = 2c/a+4, where a is the 
number of species at one site, 4, the species in another, 
and cis the number of species common to both sites. 
In addition, a turnover index (7) measured assem- 


18 THE CANADIAN FIELD-NATURALIST 


blage persistence in each site. 7 = (C+£)AS,+5,), 
where Cand £ are the number of species in respec- 
tive collections at sampling site 5, and 5, (Przybylski 
1994). Persistence (/), an indicator of stochastic 
assemblages = 1-7! All indices range from -1 to +1; a 
high value of Cc indicates a large number of species 
shared between sites. A high value of 7 indicates high 
persistence and low fish species turnover. Common 
fishes were deemed species whose frequency of occur- 
rence was >50%; uncommon occurred on 30-50% of 
sampling occasions, and rare fishes comprised the 
remainder. 

All samples were processed in the laboratory with- 
in three months of collection. Specimens were placed 
in vials containing a 12:1 solution of 80% ethanol 
and glycerin, and stored in the dark at 22-25°C. Mea- 
surements of total length (TL, mm) were made on 
most-abundant fishes and those of special interest, 
such as rare taxa or recent invader. A Wild MS dis- 
secting microscope was used for length determinations, 
accurate to 0.1 mm for fish <25 mm and 0.5 mm for 
larger specimens. For each sampling date, 10-66 speci- 
mens (average = 26) of most-abundant species were 
measured. Field records included meteorological ob- 
servations, water transparency (Secchi depth), water 
temperature, conductivity, and occasionally, pH and 
alkalinity. Environmental events, such as flooding or 
herbicide application on crops several days before 
sampling, may have affected the abundance of fishes. 
Thus, stream flow data on the sampling date and the 
previous two days were examined for possible con- 
cordance between flow volume and the total number 
of fishes found at each site. 


Results 
Environmental characteristics 

In 1994, water temperature averaged 22°C between 
June and early November, conductivity averaged 434 
uS/cem (range = 340-520 uS/cm), and Secchi disc 
0.15 to 0.50 m (mean = 0.26 + 0.13 m). In 1995, water 
temperature was maximum (29.0°C) at L in early Au- 
gust, 25.0°C at M in mid-August, and 29.5°C at U in 
mid-June (Figure 2). Conductivity was highest and 
most variable at M, where readings of 1120 and 1800 
uS/cm were recorded in late June and September, 
respectively. These values coincided with some of 
the lowest flow volumes in the river (0.012 and 0.022 
m?/d). Conductivity was lower, and transparency high- 
er at L than upstream, partly due to influx from the 
Detroit River. Eurasian milfoil first appeared in mid- 
May, and cattail, bulrush, Common Reed, American 
Lotus and Purple Loosestrife, Zy*rwm salicaria, were 
abundant in late July. Prolonged drought was respon- 
sible for a series of isolated pools (depth, 0.1-0.3 m) 
that developed at U in late June-early July. Spearman’s 
rank correlation (r,) analyses indicated total fish abun- 
dance and four environmental variables on sampling 
dates were not correlated (Table 1), nor were flow 


Vol. 119 


volume and abundance correlated on both days prior 
to sampling (r, = -0.02 to 0.20). 


Overall relative abundance and occurrence 

All reproductive guilds in Canada (Balon 1975) were 
represented in the age 0 assemblage of 42 species, 12 
families (Table 2), and total collection of 24 544 fishes. 
Clupeids (67%), dominated by Gizzard Shad, were 
the most abundant species in both 1994 and 1995; 
the remainder of the catch consisted mainly of cen- 
trarchids (13%), an atherinid (Brook Silverside, 6%) 
and 14 cyprinid species (6%). Bowfin, Spotfin Shiner, 
Banded Killifish, Blackside Darter, Tadpole Madtom, 
and Round Goby were collected only as age I* fishes. 
Several non-indigenous species (see Table 2 for sci- 
entific names), classified “Vulnerable” (Campbell 2001), 
were also found. 

Twelve common species were recorded, although 
only three contributed >5% to total catch (Table 2). 
Common Carp and Orangespotted Sunfish were the 
only common fishes among seven introduced spe- 
cies, including Alewife, Bigmouth Buffalo, Goldfish, 
White Perch and Round Goby. Maximum fish abun- 
dance occurred at all sites on the same date (23 June) 
and in similar environmental conditions in 1994 and 
1995 (Figure 3): low, declining flow (0.01-0.05 m?/d), 
low transparency, high conductivity, medium-high 
water temperature (23.5 to 29.5°C), and in L, surface 
emergence of American Lotus. 

Gizzard Shad was the most abundant age 0 fish at 
L and M, and centrarchids at U, where Orangespot- 
ted Sunfish alone represented 68.3% of total catch at 
that site. Bluntnose Minnow, Orangespotted Sunfish, 
and Quillback were the only species more abundant in 
U than downstream. White Suckers were exclusive to 
M, whereas Goldfish were caught only at U (Table 2). 

Frequency of occurrence of all taxa averaged 35% 
in L, compared with 28% and 22% in M and U, res- 
pectively. Most species, including several listed as 
“vulnerable” in Canada: Bigmouth Buffalo, Pugnose 
Shiner, Pugnose Minnow, Central Stoneroller, and 
Longear Sunfish, were caught sporadically and in low 
number. Between-site species turnover of age 0 fishes 
differed according to sampling dates, ranging from 0.9 
in early August at L and M, to 1.0 (complete lack of 
persistence) at L and U in early June. 

Overall species richness increased by a factor of 
2.2 from upper to lower reach. Assemblages of age 0 
fishes were similar taxonomically at all sites only in 
the first few weeks in June (Cc = 0.33 to 0.46). Gen- 
erally, lowest persistence, or highest turnover, also 
took place in mid-June. Highest coefficient of com- 
munity (0.5) was recorded in mid-June, when many 
taxa intermingled at all sites. The middle site and U 
usually shared fewer species than M and L. Fish co- 
occurrence, both between L and M and L and U, 
were minimum in early August (Cc = 0.13 and 0.08, 
respectively), and M and U in early July (0.07). 


2005 


Water temperature (C) 


Mean 3-day discharge (m?/d) 


LESLIE AND TIMMINS: AGE O FISHES IN RIVER CANARD 19 


—™ Lower —*- Middie —*- Upper —-— Lower 1994 


L 1994 


2000 
Bee LeU YM oe L 1994 

= ad 
E 1500 a 
7) 
= 
2 1000- 
o 
~ 
ao) 
5 
8 500 


Secchi depth (m) 


FIGURE 2. Environmental variables for River Canard: water temperature at three sites (upper), mean 3-day discharge 
(middle), and conductivity and Secchi depth (transparency) (lower graph). 


Fish Seasonality 
Upper reach 

Abundance and frequency of occurrence of age 0 
species (17 taxa) were consistently low (Figure 3). A 
single Central Mudminnow (18 mm TL) was the only 
age () fish found in late May. Quillback (15.9 + 0.8 mm; 
N = 42) next appeared (early June), but was not record- 


ed after late June. Johnny Darters (9-11 mm), which 
first appeared in early June, contributed 7.4% to the 
total catch (1839 fishes). Small numbers of recently 
hatched Common Carp (5-7 mm) were also caught in 
early June, but were not found after mid-July. No 
adult Gizzard Shad were caught on any date. However, 
Shad larvae were found sporadically until early Sep- 


20 THE CANADIAN FIELD-NATURALIST 


TABLE |. Spearman’s rank correlation coefficients for over- 
all fish abundance relative to environmental variables in River 
Canard. Upper row denotes 1994 values. Number of meas- 
urements in parentheses. 


Variable Lower reach Middle site Upper reach 
Conductivity 0.539.(12) 

0.154 (9) 0.212 (10) 0.381 (8) 
Secchi depth 0.150 (10) 

0.142 (9) -0.433 (10) 0.226 (8) 
Temperature O525 (12) 

-0.233 (9) 0.422 (10)  -0.536 (8) 
Flow volume -0.119 (12) 

-0.300 (9) -0.139(10)  -0.429 (9) 


tember, when the largest specimen (94 mm) was col- 
lected. As flow volume decreased to 0.01 m?/d in late 
June, an isolated pool (~350 m7) developed at the sam- 
pling site. Eight species and 81% of the total collection 
were caught in this pool on 23 June. Orangespotted 
Sunfish was the sole recurring adult fish. 


900 


Lower 1994 


600 


300 


Lower 1995 


500-4 


Total fishes collected 


Vol. 119 


Middle site 

Fishes represented 18 species and 7 reproductive 
guilds at the middle site. Non-guarding phytophils 
(Common Carp, Bigmouth Buffalo) and guarding nest- 
spawners, such as psammophils Bluntnose Minnows, 
Black Crappie, numerically dominated the assemblage. 
Highest catch and number of species were observed 
in late June (Figure 3). Logperch were first caught in 
mid-May and Gizzard Shad, Yellow Perch, White Suck- 
er, and Quillback in early June. Common Carp (9.4 + 
2.6 mm; N = 23), Bluegill, and Emerald Shiner next 
appeared in late June. Gizzard Shad, which represent- 
ed 78.5% of the total collection (4744 fishes), was the 
only highly persistent species (Table 3). Although var- 
ious age 0 and 1+ fishes co-occurred in early spring 
and autumn, they were usually temporally separate dur- 
ing the early summer nursery period. Emerald Shiner 
was the only adult fish caught repeatedly. 


Lower reach 

Although sampling in 1994 began in late June, 12 
taxa were found that were not caught in 1995, when 
collections began in April. However, these fishes con- 


Number of taxa 


M J J A S 


FIGURE 3. Seasonal total catch (bar) and number (line) of age 0 fishes found in River Canard, 1994-1995. 


2005 


TABLE 2. Age 0 fish (0) species in River Canard lower (L), middle (M), and upper (U) sites, 1994-1995. 1+ indicates occur- 


LESLIE AND TIMMINS: AGE O FISHES IN RIVER CANARD 


21 


rence of juveniles >1 year old, or adults. f,= relative overall occurrence: C common, U uncommon, R rare. Guild = repro- 


ductive classification (Balon 1975): Ph phytophil, Lp lithopelagophil, Pl phytolithophil, Ps psammophil, Pe pelagophil, Sp 
speleophil, P polyphil. Total catch 24544. TL (mm) indicates smallest »zeasured specimen of each species. Species status: N 


Native, I introduced, V “vulnerable”. Species names follow Mandrak and Crossman (1992). 


Species 


Lepisosteus OSSCUS 
Amia calva 

Dorosoma cepedianum 
Alosa pseudoharengus 
Evsox luctus 

Umbra limi 


Moxostoma macrolepidotum 


Moxostoma sp. 
Catostomus commersont 
Ictiobus cyprinellus 
Carpiodes cyprinus 
Notropis atherinoides 
Notropis hudsonius 
Notropis volucellus 
Notropts stramineus 
Luxilus cornutus 
Notropis anogenus 
Opsopoeodus emiliae 
Notemigonus crysoleucas 
Cyprinus carpio 
Carassius auratus 
Cyprinella spiloptera 
Campostoma anomalum 
Pimephales notatus 
Pimephales promelas 
Ameturus nebulosus 
Ictalurus punctatus 
Noturus gyrinus 
Fundulus diaphanus 
Morone chrysops 
Morone americana 
Lepomis gibbosus 
Lepomis megatlotis 
Lepomis cyanellus 
Lepomis humilis 
Lepomis macrochirus 
Ambloplites rupestris 
Micropterus salmoides 
Micropterus dolomieu 
Pomoxis nigromaculatus 
Pomoxts annularis 
Perca flavescens 
Etheostoma nigrum 
Percina caprodes 
Percina maculata 
Labidesthes sicculus 
Aplodinotus grunniens 
Neogobius melanostomus 


Common name 


Longnose Gar 
Bowfin 

Gizzard Shad 
Alewife 

Northern Pike 
Central Mudminnow 
Shorthead Redhorse 
Redhorse sp. 
White Sucker 
Bigmouth Buffalo 
Quillback 

Emerald Shiner 
Spottail Shiner 
Mimic Shiner 
Sand Shiner 
Common Shiner 
Pugnose Shiner 
Pugnose Minnow 
Golden Shiner 
Common Carp 
Goldfish 

Spotfin Shiner 
Central Stoneroller 
Bluntnose Minnow 
Fathead Minnow 
Brown Bullhead 
Channel Catfish 
Tadpole Madtom 
Banded Killifish 
White Bass 

White Perch 
Pumpkinseed 
Longear Sunfish 
Green Sunfish 


Orangespotted Sunfish 


Bluegill 

Rock Bass 
Largemouth Bass 
Smallmouth Bass 
Black Crappie 
White Crappie 
Yellow Perch 
Johnny Darter 
Logperch 
Blackside Darter 
Brook Silverside 
Freshwater Drum 
Round Goby 


Status 


ZZALBAALZLZALZ 


< 


ZZ, 2 2, ZZ 


eS 
<2 


ee Pee re ee eee ee oe dee 


tributed just 3% to the overall catch. Conversely, Log- 
perch, Smallmouth Bass, and Longnose Gar, found in 
1995 were not caught in 1994; these fishes were caught 
in small number, usually on a single date. Gizzard Shad 
(25.4%), Brook Silverside (21.9%), Orangespotted 
Sunfish (19.2%), and Bluegill (13.9% of total catch) 


L 
0 


were most abundant of 36 species and 3055 specimens 
collected in 1994. Frequency of occurrence of these 


M 


O, 1+ 


U 


Guild 


f 


(9) 


AAIAAAIAAXZINAAZA AINA A AAA AARZAAINANANAAAAARDAIINNCAARAARAARAAADA 


Catch (%) 


66.6 


8.4 
bee) 


6.3 


TE 
80 


four taxa was also highest (f, = 0.8-1.0) throughout 


the sampling period. Orangespotted Sunfish and Spot- 
tail Shiners were the only adult fishes collected con- 


sistently. 


22 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE 3. Percent contribution, seasonality, and frequency of occurrence (parentheses) of dominant fishes at three collection 
sites in River Canard, 1994-1995. 


Species Lower Mid Upper 
1994 1995 
Gizzard Shad 25.4 June-November 80.7 June-October 78.5 June-September 6.2 June, August, 
(1.0) (0.9) (0.9) September (0.3) 
Emerald Shiner 5.4 June-November 0.6 June-August (0.4) June-July (0.3) 
(0.2) 
Common Carp 0.2. June-July (0.3) 0.5 June-August (0.4) June-July (0.4) June (0.2) 
Bluntnose Minnow 1.2 June-August 0.2 June-July, June (0.1) 13.7. June-August 
(0.6) September (0.5) (0.6) 
Orangespotted Sunfish 19.2 June-November 0.6 June-August June-July 68.3 June-September 
(0.9) (0.6) (0.2) (0.6) 
Bluegill 13.9 June-November 3.1 June-September June-July 1.8 June, September 
(0.8) (0.8) (0.4) (0.2) 
Brook Silverside 21.9 June-November 10.1 June-September July (0.1) 
(0.9) (0.6) 
Total catch 2671 8452 1089 1532 
TABLE 4. Comparitive growth in length of eight species of age-0 fishes in the Great Lakes basin. 
Species Mid-month TL (mm) 
System Year August September Reference 
Gizzard Shad Lotic 1994 62.2 (42) 80.2 (66) Present study 
1995 69.7 (26) 77.8 (19) 
Lotic 1994 48.5 (23) 7233 27) Leslie et al. 1999 
Lentic 1994 45.7 (£5) 58.2 (26) Leslie and Timmins 1998b 
Lentic 1990 51.9 (20) 74.0 (18) Leslie and Timmins 1998b 
Lacustrine 1983 32.002) Leslie and Timmins 1993 
Brook Silverside Lotic 1994 42.6 (25) Present study 
Lotic 1994 44.6 (24) Leslie et al. 1999 
Lentic 1994 46.0 (23) Leslie and Timmins 1998b 
Lacustrine 1983 43.0 (59) Leslie and Timmins 1993 
Emerald Shiner Lotic 1994 33.9135) 43.5 (14) Present study 
Lotic 1994 34.7 (43) 41.5 (26) Leslie et al. 1999 
Lotic 1994 44.9 (39) 48.2 (32) Leslie et al. 1999 
Lacustrine 1994 42.4 (21) Leslie and Timmins 1998c 
Bluntnose Minnow Lotic 1994 26.0 (15) Present study 
1995 29.5 (24) 
Lotic 1994 45.3 (8) 50.2 (20) Leslie et al. 1999 
Lacustrine 1977 36.4 (72) 51.3 (79) Keast and Eadie 1984 
Lacustrine 1998 29.7 (55) Leslie and Timmins 2002 
Orangespotted Sunfish Lotic 1994 38.1 (23) Present study 
1995 34.6 (15) 
Lentic 1990 36:5 (22) Leslie and Timmins 1998b 
Bluegill Lotic 1994 30.6 (37) 36.9 (26) Present study 
1995 31-0115) 34.4 (48) 
Lotic 1994 33.367) Leslie et al. 1999 
Lacustrine 1977 16.4 (56) 26:7 G5) Keast and Eadie 1984 
Lentic 1990 28.7 (19) Leslie and Timmins 1998b 
Black Crappie Lotic 1994 56.7 (26) 59.9 (15) Present study 
1995 62.2 (29) 
Lotic 1994 5525652) 6L.2:17) Leslie et al. 1999 
Largemouth Bass Lotic 1994 91.9 (10) Present study 
1995 72.6 (10) 
Lacustrine 1977 48.6 (89) Keast and Eadie 1984 


2005 LESLIE AND TIMMINS: AGE O FISHES IN RIVER CANARD 23 
i Larvae Juveniles 
40 1 x8 rice Sliven} sons al Joes 
35 a” ele: @Dc OLm ALh XLs. 
a pee wes 
cn ge? Bi qrinr attr spulfmentt ats 
45 — ae Ox OO» ¢ 
10 - @ en A x od e 
5 | } 
0 +— — ——— ee 
0 20 40 60 80 100 
Mean TL (mm) 


FIGURE 4. Coefficient of variation in mean total lengths of Gizzard Shad (Pc), Bluegill (Z77), Orangespotted Sunfish (ZA), 
and Brook Silverside (Zs) in River Canard, Ontario. Vertical line delineates larvae (<25mm) and age 0 juveniles. 


Total catch in 1995 (8771 fishes, 26 species) was 
dominated by Gizzard Shad (80.7%), Brook Silverside 
(10.1%) and Bluegill (3.1%). No fish larvae were found 
in April or May. Largest catches of Gizzard Shad (4- 
5 mm), Orangespotted Sunfish and Bluegill were made 
in late June. Spottail Shiner, Bluegill, Black Crappie, 
Brook Silverside, Bluntnose Minnow, and Largemouth 
Bass were considered “residents” based on high per- 
sistence. As in 1994, adult Orangespotted Sunfish and 
Spottail Shiner were collected on most dates. 

Plankton net hauls in open water collected 5280 age 
0 fishes (10 species) in mid-June, 1995. None of these 
species was unique to the assemblage collected with 
beach seines. However, because Freshwater Drum 
spawn in open water where semi-buoyant eggs drift 
with currents, the plankton net collected more eggs 
and free embryos (3.7 + 0.4 mm; N = 13) than did the 
beach seine. Total catch was dominated by Gizzard 
Shad (97.3%), Common Carp (1.0%) and Freshwater 
Drum (0.6%). Mean fish density varied widely among 
hauls and dates. For example, mean densities ranged 
from 4201 to 6390/100 m? and 1199 to 2419/100 m? 
on 11 June and 12 June, respectively. 


Growth of most-abundant fishes 

Lengths near the end of first year growth differed 
according to species. Bluntnose Minnow and Brook 
Silverside were the only fishes that nearly attained adult 
length in late summer. Gizzard Shad, Largemouth Bass, 
and Black Crappie are among the largest fishes in the 
assemblage (Table 4). These species achieved 12 to 
25 times their length at hatch, whereas small-bodied 
fishes (e.g., cyprinids, Brook Silverside) increased 6 
to 8-fold hatch size. Coefficient of variation (CV) in 
mean total length of four most-abundant species was 
highest in larvae (<25 mm) and tended to decrease with 


size (Figure 4). In general, highest variability in species’ 
lengths coincided with overall peak abundance in mid- 
June to mid-July. 


Discussion 

The assemblage of age 0 fishes (42 taxa) may be 
considered a cyprinid-centrarchid-clupeid-atherinid 
complex numerically representing 91% of the catch 
(Table 2) and 63% of a taxocene strongly dominated 
by a planktivore, Gizzard Shad. Lithophils (non-guard- 
ing and guarding) and phytophils otherwise predomi- 
nate in constant turbid water and shoreline with limited 
diversification. Larvae of these guilds are initially pho- 
tophobic (Mann 1996) and in this respect, perhaps tol- 
erant of turbid conditions. However, the presence of 
most species in River Canard belies spawning require- 
ments for gravel, macrophytes, and clear water. Orange- 
spotted Sunfish selects shallow, silt-laden water (Noltie 
and Beletz 1984) and is an appropriate indicator spe- 
cies, contrary to Bluegill, which usually chooses vege- 
tated shallows in clear water (Trautman 1981). 

Similarly, Emerald Shiner, a pelagophil, was com- 
mon and abundant at the middle site and in the lower 
reach. It was also found, albeit in low numbers, in adja- 
cent watersheds, shore areas, and streams in south-east 
Lake St. Clair and western Lake Erie (Noltie and Beletz 
1984; Leslie and Timmins 1998a,b,c; Leslie et al. 1999). 
Undoubtedly, this is a plastic species with considerable 
ecological amplitude, as is the Bluntnose Minnow. 
Brook Silverside, another common taxon, usually 
spawns on emergent or floating leaf plants, hence its 
absence in the upper reach and presence in the lower 
reach. The assemblage in the lower reach represented 
many ecological guilds dominated by persistent fishes, 
such as Gizzard Shad, Brook Silverside, Emerald Shin- 


24 THE CANADIAN FIELD-NATURALIST 


er, Orangespotted Sunfish, and Bluegill. Gizzard Shad 
spawned in the upper reach and undoubtedly further 
upstream in connecting minor drainage systems, where 
larvae are found in vast number (Leslie and Timmins 
1990). 

Most fishes expanded their ecological limits with 
growth and development. Species turnover was typi- 
cally high and relatively few taxa occurred simulta- 
neously at the three sites. These findings concur with 
those in similar studies in adjacent watersheds of the 
St. Clair and western Lake Erie flatlands (Leslie and 
Timmins 1990, 1998a; Leslie et al. 1999). Commonly, 
species richness and abundance are highly correlated 
with macrophyte diversity and shallow marginal habi- 
tat that provides refuge against predators (Eadie and 
Keast 1984; Petering and Johnson 1991; Copp and 
Mann 1993; Mann 1996; Duncan et al. 2001). Sinuous 
and sluggish, River Canard is devoid of vegetated 
backwaters that enhance diversity, spawning and nurs- 
ery habitats, and food availability (Oberdorff et al. 
1993). Nevertheless, Pugnose Minnow, Pugnose Shin- 
er, and Central Stoneroller, sensitive to turbid waters, 
tolerated adverse conditions, at least temporarily. 

Environmental variables and species numbers were 
not related with seasonal abundance (Table |; Figure 
3). Number and abundance of age 0 fishes in the lower 
reach were likely augmented by continuous larval drift, 
individuals purged from upstream during spates, and 
migrants from connecting drainage ditches and the 
Detroit River. Although River Canard appears to be a 
harsh environment for fishes during early ontogeny, 
large numbers of taxa have adapted to, if not thrived, in 
this highly perturbed system. Abiotic and biotic fac- 
tors influence rate of growth during early development, 
and hence, recruitment. Total length attained by re- 
cruits near end of summer may well reflect these fac- 
tors, providing sampling does not seriously bias size 
distributions of any given species (Bayley and DowI- 
ing 1993; Leslie and Timmins 1994). 

With few exceptions, dominants were much the same 
length as those recorded elsewhere in the ecoregion 
(Table 4). For example, Gizzard Shad and Emerald 
Shiner reached autumnal lengths comparable to those 
in turbid streams and bays but less than in waters of 
higher quality, transparency and productivity, such as 
the Detroit River (Leslie et al. 1999). Orangespotted 
Sunfish were slightly longer in River Canard than 
reported for specimens in lowland Ohio streams (Traut- 
man 1981), whereas Bluntnose Minnows were small- 
er than counterparts in the St. Clair region. Similarly, 
Bluegill and Brook Silverside attained lengths similar 
to those of species elsewhere in the lower Great Lakes 
basin (Trautman 1981; Leslie and Timmins 1990; 
1998a,b; Leslie et al. 1999). Size range relative to mean 
total length of cyprinids and other small species was 
usually much lower for age 0 juveniles than for larvae, 
as was overall coefficient of variation in length. Sam- 
pling bias, predation, and natural mortality of small- 


Vol. 119 


est fish may effect a reduction in variability in length 
as growth reaches a plateau. 

The diverse age 0 fish taxocene in River Canard is 
represented by all reproductive guilds in Canada. Yet, 
according to descriptions in Scott and Crossman (1973) 
and Trautman (1981), fewer than 25% of species listed 
in Table 2 normally utilise environmental conditions 
that prevail in the river. Apparently, assignment of cer- 
tain species to a specific reproductive or ecological guild 
requires revision for Great Lakes as well as European 
freshwater fishes (Mann 1996). Perhaps these realities 
reflect our limited knowledge of fish ecology in low- 
land streams. 


Acknowledgments 

We thank Susan Saunders of Water Surveys of Can- 
ada, Burlington, for flow data for River Canard, R.E. 
Dermott for assistance in the field, and several anony- 
mous reviewers. 


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Received 20 May 2003 
Accepted 22 November 2004 


Charophytes of Insular Newfoundland Il: Chara evoluta and Chara 
CANESCENS 


HENRY MANN and E. M. V. NAMBUDIRI 


Environmental Science/Biology, Sir Wilfred Grenfell College, Memorial University of Newfoundland, Corner Brook, New- 
foundland and Labrador A2H 6P9 Canada 


Mann, Henry, and E. M. V. Nambudiri. 2005. Charophytes of Insular Newfoundland II: Chara evoluta and Chara canescens. 
Canadian Field-Naturalist 119(1): 26-37. 


The rare Newfoundland disjuncts Chara evoluta and Chara canescens are compared morphologically, ecologically, and geographi- 
cally. Although very similar morphologically, they can readily be distinguished by their monoecious/dioecious nature, cortical 
cell features and coronula size. Ecological differences are more subtle and difficult to quantify. In Newfoundland the two 
species are restricted to coastal saline lagoons in the southwest region of the Island, with C. evo/w/a exhibiting an association 
with Chara aspera. The flora and fauna of these unique habitats are poorly documented even though it is known that they 
are associated with other rare species including the Piping Plover and the Banded Killifish. Consideration of some degree of 
protection as special habitats within the provincial ecological reserve strategy is suggested. An updated key to all species of 
charophytes known to inhabit Newfoundland and Labrador is included, and a brief overview of their ecological distributions 


is provided. 


Key Words: charophytes, Characeae, Chara evoluta, Chara canescens, Chara aspera, coastal lagoons, Newfoundland. 


The first survey of the charophytes of Newfoundland 
(Mann 1989) described ten taxa, three of V7e//a and 
seven of Chara. Distribution maps and a key were 
provided. Subsequently 70/ypel/la glomerata (Desv.) 
Leonh. was reported from the Great Northern Penin- 
sula (Mann 1994a). In 1995 Chara evoluta T. F. Allen 
was discovered as well as a second site for Chara 
canescens Desv. & Lois. in Lois. and these were added 
to the list for Insular Newfoundland (Mann 1998, 
2000*; Mann et al. 1999). A search of saline coastal 
lagoons and estuaries on the southwest coast of New- 
foundland identified several more sites for C. evo/ufa, 
but no more for C. canescens. Information, including 
morphological, taxonomic, geographical and ecolog- 
ical, has never been published for this disjunct popu- 
lation of C. evo/uta in Newfoundland. Because the two 
species are so similar morphologically and ecologi- 
cally, and because both C. evo/uta and C. canescens 
are considered rare species in Newfoundland, eastern 
Canada, and C. evo/ufa in eastern North America, the 
two will be treated together in a comparative way. 
They also need to be distinguished and reported as 
separate taxa, a problem which arose since the publi- 
cation of Wood’s (1965) monograph where they were 
merged under C. canescens, a concept which is no 
longer defensible (Proctor 1980). Due to the disjunct 
nature of Newfoundland populations it is important 
to determine whether local populations exhibit any fea- 
tures differing from those elsewhere in North America. 
This is also necessary for future studies comparing 
similar Asian taxa. T. F. Allen’s (1882) original des- 
cription of C. evo/uta and those of Robinson (1906), 
and Wood’s chosen lectotype in Wood and Imahori 
(1964) are the basic sources for this species supple- 
mented with North American specimens on loan from 


the New York Botanical Garden (NY). It may be as- 
sumed that features not here described or elaborated 
upon are consistent with those in the above reports. 

A new key is prepared incorporating species not 
included in the original work (Mann 1989) and based 
on current concepts of charophyte taxonomy and 
nomenclature. 

Material of C. evoluta and C. canescens is housed in 
the Sir Wilfred Grenfell College Herbarium (SWGC) 
as liquid preserved specimens and herbarium sheets. 
Voucher specimens have been deposited with the 
Newfoundland Museum (NFM), the Canadian Muse- 
um of Nature, Ottawa (CANA), and the Phycological 
Herbarium of Memorial University (NFLD), (C. evo- 
/uta, Mann numbers 228, 282; C. canescens, Mann 
number 229). Herbarium acronyms are as in Holm- 
gren et al. (1990), except for the Sir Wilfred Grenfell 
College (SWGC) which is not yet listed. Vascular plant 
nomenclature follows Crow and Helquist (2000), that 
of invertebrate groups, Barnes (1994), and fishes, Scott 
and Crossman (1973). Methods for gametangial meas- 
urements and illustrations are consistent with those 
described in Mann (1994b). Spore preparation for the 
SEM followed the method of John and Moore (1987). 


Study Area and Habitat 

The study area where C. evo/uta and C. canescens 
are located includes south-western Newfoundland ex- 
tending from Port aux Basques in the south and north- 
ward to the Port au Port Peninsula (Figure 1). All sites 
are coastal saline lagoons or salt ponds, sometimes 
designated as “barachois” locally. The southernmost 
cluster of five lagoons can be located on Canadian 
Topographic Map 11-0/11, “Port aux Basques’. Cod- 
roy Estuary Pond has UTM map co-ordinates of 314E 


26 


2005 


MANN AND NAMBUDIRI: CHAROPHYTES OF INSULAR NEWFOUNDLAND II ZI 


FIGURE |. Location of collection sites in Insular Newfoundland. 


A. Study area of southwestern Newfoundland inset. 


B. Coastal Newfoundland from Port aux Basques to the Port au Port Peninsula: (a) Gravels Pond, (b) St. George’s Pond, 


(c) Codroy Estuary Pond, (d) Port aux Basques. 


C. Isthmus of the Port au Port Peninsula: (a) Gravels Pond. Dashed lines indicate roads. 
D. St. George’s Bay: (a) Main Gut, (b) Estuary, (c) St. George’s Pond. Dashed lines indicate roads and a railway 


embankment. 


E. Coastal Newfoundland from Cape Ray to Grand Bay: (a) Cape Ray, (b) Little Barachois, (c) Osmond Pond, (d) 
Big Barachois, e. Rocky Barachois, (f) Salt Water Pond, (g) Second Pond, (h) First Pond, (i) Grand Bay. 


and 018N on map 1 1-0/14, “Codroy”. The more north- 
em Gravels Pond is indicated on map 12-B/10, “Steph- 
enville”, and the St. George’s Lagoon on map 12-B/8, 
“Main Gut”. 

The southern five lagoons from Grand Bay to Cape 
Ray Cove are formed behind sandy beach and dune 
deposits. All have breachways to the ocean, but the 
channels are not active during most of the year and no 


seawater exchange is apparent normally, except for 
First Pond where exchange may occur more often. 
Outflow occurs during spring runoff, but there is evi- 
dence that some saltwater influx occurs to maintain 
their brackish nature. All of these ponds drain fresh- 
water from inland peatlands and heath barrens. The 
small Codroy Estuary Pond is in the floodplain of the 
estuary, but is isolated from estuarine water except 


28 THE CANADIAN FIELD-NATURALIST 


during spring flooding. St. George’s Lagoon is sepa- 
rated from the ocean by a sand barrier of 100 meters 
in width. Originally connected to the estuary, it is now 
separated by highway and railway embankments, but 
still receives some influx via culverts where saline 
estuarine water intermixes with the freshwaters of 
Little River draining the interior peatlands. Gravels 
Pond in the narrow isthmus of the Port au Port Penin- 
sula is separated on either side from the ocean by a 
narrow cobble/shingle barrier. 

Coastal lagoons in this study are non-tidal, but only 
semi-isolated systems receiving some periodic seawa- 
ter influx. As such, the salinity of their waters varies 
with the seasons and is also subject to rapid fluctua- 
tions because of heavy rains, storm surges, salt spray, 
and perhaps some percolation through the barrier as 
possibly occurs in Gravels Pond. Because of irregular 
sporadic episodes of varying intensity they are habitats 
extremely difficult to characterize and compare other 
than through intensive long-term monitoring (Barnes 
1980). In 1995 Gravels Pond ranged in specific con- 
ductance (umhos = yu siemens) from 10 000 in July, 
15 200 in August, to 22 200 in September. According 
to the salinity scale of Cowardin et al. (1979), Grav- 
els Pond is in the brackish mesohaline category. First 
Pond, Second Pond, and Saltwater Pond exhibited 
single readings of 26 400, 18 000, and 27 000 umhos 
respectively in July 1999, also categorizing them as 
mesohaline. St. George’s Lagoon had the lowest read- 
ings ranging from 400 to 7500 umhos characterizing 
it as fresh to oligohaline. Internally, lagoons are often 
not uniform with regard to salinity, possessing zones 
depending on the location and volume of freshwater 
entry as well as periodic marine influx (Barnes 1980). 

The Newfoundland lagoons which are separated 
from the ocean by a sandy barrier have extensive shal- 
low sandy-bottomed areas on their seaward sides due 
to windblown sand drift from the outer beaches and 
gradually deepen to several meters on their landward 
sides. This produces an unstable sandy shifting sub- 
strate subject to wave action where vascular vegeta- 
tion is sparse or absent, but to some extent success- 
fully colonized by charophytes. On these shallow flats, 
charophytes form open scattered stands where indi- 
vidual plants tend to be tiny, only a few centimeters, 
and deeply embedded in the sand substrate. In deeper 
areas or sheltered locations with more stable substrates, 
denser beds of larger charophytes and beds of vascu- 
lars occur. 

Aquatic vasculars largely consist of broad-ranging 
and saline tolerant species including Sago Pondweed 
(Potamogeton pectinatus L. = Stuckenta pectinata \L.) 
Borner), Widgeon-grass (Ruppia maritima L.), and 
Horned Pondweed (Zannichellia palustris L.). In zones 
of lower salinity Clasping-leaved Pondweed (/ora- 
mogeton perfoliatus L.) and Mare’s-tail (A/ippuris vul- 
garis L.) are occasionally found. Eelgrass (Zostera 
marina L.) is absent from these charophyte dominated 
lagoons, but is common and forms extensive stands 


Vol. 119 


in tidal lagoons which also occur in the area (e.g., Big 
Barachois, Figure 1). Charophytes have not been locat- 
ed in the tidal lagoons, but potentially may occur near 
freshwater inlets. 

Although the aquatic fauna was not systematically 
sampled, the following organisms were commonly 
noted: amphipods, water boatman, snails, opossum 
shrimps, aquatic beatles, and a variety of insect larvae 
forms. Two fish species were encountered. The Thee- 
spine Stickleback (Gasterosteus aculeatus L.) is com- 
mon in all lagoons amongst the charophyes. As well, 
in First Pond the Banded Killifish (4zdulus diaphanus 
Leseur) was commonly in the charophyte beds. 

Species diversity decreases with increased salinity 
in inland waters (Cole 1994), but in addition, coastal 
lagoons often present a widely fluctuating ionic envi- 
ronment which cannot be tolerated by most marine 
organisms as well as most freshwater derived inland 
forms. Biodiversity, therefore, tends to be low and 
lagoons are inhabited by organisms that have a broad 
tolerance to fluctuating conditions. Lagoons also tend 
to contain a narrow range of niches and mainly favour 
a suite of organisms adapted to shallow, soft sediment 
habitats (Barnes 1994; 1999). Nevertheless, like estu- 
aries, lagoons are highly productive and important 
feeding and migratory sites for waterfowl] and shore- 
birds (Nelson-Smith 1977; Barnes 1980; Gillespie et 
al. 1991). 


Morphology 

Newfoundland Chara evoluta produces a cluster 
of axes arising from an enlarged basal node anchored 
in the substrate by rhizoids. Axes up to 23 cm. in 
length and 520 um in diameter have been observed 
from depths of one to several meters, but in very 
shallow waters plants of only 2 to 3 cm. in length can 
be commonly found. Main axes are basically hap- 
lostichous, that is, having only vertical rows of corti- 
cal cells equal in number to the number of branchlets 
in a whorl. These cortical rows of cells are all primary 
rows with all rows containing spine cell clusters. Al- 
though the haplostichous condition may be quite reg- 
ular, invariably in this species small secondary cells 
can be found along the axis between the primary cells 
producing an imperfect haplostichous condition and 
showing a tendency towards the diplostichous condi- 
tion (Figure 2A). Spine cells are produced at the nodes 
of the cortical cells, sometimes singly, but more often 
in clusters of two or three. Characteristically, at least 
one spine cell at a node will be long, up to twice the 
diameter of the axis, whereas the other one or two are 
shorter, often being quite tiny and rudimentary (Fig- 
ure 2A, 2D). Two rows of stipulodes are produced at 
the base of the branchlets, the cells of the upper row 
being slightly longer than the lower (Figure 2A, 2B). 

Whorls of 9 to 10 branchlets occur at intervals along 
the axis. Branchlets are corticated with whorls of bract 
cells at the nodes (Figure 2B). A small ecorticate end 
cell arises from the terminal whorl. In some collections 


2005 


OE 


ei 
| 
} 


MANN AND NAMBUDIRI: CHAROPHYTES OF INSULAR NEWFOUNDLAND II 29 


FiGuRE 2. Morphological features of C. evo/uta and C. canescens. 


C. evoluta branchlet. Scale bar = 1000 um. 


aK) C2. 


an elongated branchlet extension occurs of 2 or 3 
ecorticate end cells (Figure 2C). Invariably these 
plants have been heavily intermingled with filamen- 
tous algal growth so this feature is probably a stress 
reaction to local biotic or abiotic conditions. This 
feature is not seen in normal vigorous material grow- 
ing without obvious competition. 

Gametangia, antheridia and oogonia are conjoined 
at the lowest 2 to 3 branchlet nodes (Figure 2B). Al- 
though the antheridia are produced below the oogo- 
nia at the nodes, in plants with very short branchlets 


C. evoluta axis - a. secondary cortical cells. Scale bar = 500 um. 


C. evoluta variation in ecorticate end cells of branchlets; unusual, ecologically induced. Scale bar = 1000 um. 
C. evoluta typical spine cell clusters. Scale bar = 500 um. 
C. canescens typical spine cell clusters. Scale bar = 1000 um. 


the internodes are too short to allow antheridia and 
oogonia to line up above each other and a superficial 
appearance of two rows is produced with antheridia 
and oogonia lateral to each other. The two bracteoles 
on either side of the oogonium are usually shorter or 
as long as the mature oogonium whereas the bract cells 
tend to be much longer. Gametangial measurements 
of C. evolura are listed in Table | and compared to 
those of the lectotype provided by Wood and Imahori 
(1964). Newfoundland measurements compare favour- 
ably with this originally described material from west- 


30 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |. Gametangial and oospore features of Chara evoluta and Chara canescens. Measurements are presented as means 


(um) with range values in parentheses. 


C. evoluta 
ce. Wood and 
evoluta Imahori 
Newfoundland 1964 
Oogonium 
length* 931 (816 — 1002) 690 — 810* 
width 474 (372 — 548) 420 — 450 
coronula height 128 (103 — 155) 90 — 105 
coronula width 249 (207 — 279) 200 — 220 
convolutions (12 — 15) (13 — 15) 
Oospore 
length 601 (517 — 661) 585 — 645 
width 338 (227 — 403) 320 — 360 
Number of ridges* (9 — 12) (10 — 12) 
fossae width* 55 (47 — 65) 51 
Antheridium 
diameter 348 (310 — 393) 300 — 330 


Cc ec; 
C: CANESCENS CANESCENS 
CANESCENS Wood and Muenscher Krause 
Newfoundland 1956 1997 
759 (620 — 878) 710 550 — 800* 
400 (341 — 455) 464 300 — 450 
71 (62 — 83) 71 50 — 80 
137 (103 — 165) 178 100 — 200 
(10 — 15) (12 — 13) (11 - 15) 
557 (444 — 671) 535 400 — 700 
299 (258 — 341) 357 200 — 400 
(10 - 14) 11 (10 — 13) 
50 (42 — 55) - - 
— — 500 — 700 


* Length measurements include coronula length except Wood and Imahori (1964) and Krause (1967) which do not. 
* Number of measurements (n) are 100 in all Nfld. features, except oospore No. of ridges (n = 50) and oospore fossae 


width (n = 30). 


ern Canada. Wood and Imahori (1964) state that the 
antheridia may be 4 scutate, but indicate with a (?) their 
uncertainty. No other reports mention this feature. All 
members of the genus C/ava at present are considered 
octoscutate with the exception of C. zey/anica Klein 
ex Willd. which is tetrascutate (Proctor et al. 1971). 
Determining the number of antheridial plates (scutes) 
in dried herbarium material is often not possible and 
scutes of liquid preserved material of this species are 
even difficult to separate clearly. Newfoundland mate- 
rial is distinctly octoscutate and we suspect this 1s equal- 
ly true for the lectotype. Characteristic of the oospore 
are the very low, almost inconspicuous ridges, and the 
claws of the cage which are almost always present at 
the base (Figure 3D, 3E). The number of ridges (striae) 
per oospore ranged from 9 to 12, but most oospores 
(72%) had 10 or 11. 

Chara canescens Desv. & Lois. in Lois. is morpho- 
logically very similar to monoecious C. evo/uta, but 
is not known to produce antheridia in North America, 
instead producing oospores parthogenetically. Besides 
the obvious absence of plants with antheridia, a num- 
ber of other morphological differences occur, although 
some being quite subtle unless one is familiar with 
both species. The best, most obvious, and most con- 
stant of these is the comparative size of the coronulas 
(Table 1, Figure 3C, 3F). Coronulas of C. evo/uta are 
almost double the size of C. canescens and there is no 
overlapping in the range values. The tips of the spiral 
cells just beneath the coronula also tend to enlarge 
considerably more at maturity in C. evo/u/a than in 
C. canescens. Detached oogonia of the two species can 
easily be distinguished by these features. Other game- 
targial measurements (Table 1) are consistently lower 
in C. canescens but not sufficiently to be taxonomically 


useful. Oospores of C. canescens tend to exhibit more 
ridges with 72% of the oospores having 11 or 12 striae. 
The striae are also more pronounced than those of C. 
evoluta (Figure 3A, 3D), a feature readily seen with 
the compound light microscope as well. Basal claws 
are normally absent, but normally present in C. evo- 
/uta. Oospore membranes of both species are indis- 
tinctly and minutely granular (Figure 3B, 3E). Com- 
parative measurements of North American C. canescens 
are provided by Wood and Muenscher (1956) from 
New York State (Table 1). European measurements 
are also given from Krause (1997) although these com- 
bine sexual dioecious and parthenogenetic forms as 
will be discussed in the taxonomic section. All three sets 
of measurements are fairly consistent for C. canescens. 

Several subtle vegetative differences also occur 
between the species. C. canescens produces a more 
perfectly uniform haplostichous axis, only rarely are 
tiny secondary cells formed between primary cortical 
rows. Up to 4 to 6 spine cells are normally produced 
in a cluster (Figure 2E) and all or most tend to be 
equally long, often twice as long as the axis diameter 
producing a very spiny appearance. On the other hand, 
C. evoluta normally produces | to 3 cells per cluster, 
one or two often much smaller (Figure 2D). 


Taxonomy 

Chara evoluta T. F. Allen was first described by T. 
F. Allen from the Red Deer Lakes and the saline 
ponds west of the Saskatchewan River (Alberta) based 
on material collected by J. Macoun (Allen 1882). 
Founded on a belief that the monoecious/dioecious 
conditions are not good indicators of species bound- 
aries, Wood (1965) combined C. evo/uta with dioe- 
cious and parthenogenetic taxa under C. canescens 


2005 


eS) 


FIGURE 


amo mw > 


MANN AND NAMBUDIRI: CHAROPHYTES OF INSULAR NEWFOUNDLAND II 


. Oogonium and oospore features of C. evo/uta and C. canescens. 
C. canescens spore (SEM). Scale bar = 120 um. 

C. canescens spore membrane (SEM). Scale bar = 30 um. 

C. canescens oogonia (Light Microscope). Scale bar = 500 um. 
C. evoluta spore (SEM). Scale bar = 100 um. 

C. evoluta spore membrane (SEM). Scale bar = 24 um. 

C. evoluta oogonia (Light Microscope). Scale bar = 500 um. 


32 THE CANADIAN FIELD-NATURALIST 


Desv. & Lois. in Lois. Subsequently it has been ade- 
quately demonstrated that monoecious and dioecious 
morphologically similar species pairs in charophytes 
are not conspecific and that Wood’s views on this 
matter are no longer tenable (Proctor 1980). Never- 
theless, this combining has led to continuing difficulty 
in distinguishing taxa in literature reports ever since, 
a matter especially troublesome for biogeographic 
purposes. 

T. F. Allen (1900) described a very similar taxon 
which he designated as Chara hirsuta from a fresh- 
water pond in California. It is a more robust plant to 
60 cm. in height with somewhat larger oospores, more 
striae, and somewhat more prominent ridges. By com- 
paring the descriptions based on the same material 
collected from Lakeside, California, by Allen (1900) 
and others (Robinson 1906; Wood 1965) it becomes 
apparent that considerable overlap exists in the mor- 
phological features of C. Azrsuta and C. evolura. \t has 
long been understood that morphological variants of 
charophyte species can often be encountered in iso- 
lated bodies of water, but that these minor ecological 
or genetic variants probably do not indicate repro- 
ductive isolation (Robinson 1906; Moore 1986). One 
report of C. Airsu/a occurs from Nebraska (Daily and 
Kiener 1956) and one from Wyoming (Daily and Porter 
1961), however, Daily and Kiener (1956) suggest that 
C. evoluta and C. hirsuta are probably conspecific. 
That view was also taken by Proctor (1990) and is ac- 
cepted here as well, although no evidence other than 
morphological is available to support this conjecture. 

A monoecious, morphologically similar taxon to 
North American C. evo/ufa has been reported from 
Asia designated as C. a/raica A.Br. = C. sibirica 
Mig. (Wood 1965; Hollerbach and Krassavina 1983). 
The C. evoluta taxon has also been reported from 
China (Han and Li 1994). In light of Proctor’s (1980) 
conjecture that most charophytes are endemic to a sin- 
gle land mass, it remains to be seen whether the North 
American and Asian taxa are conspecific. To our know- 
ledge no such comparative studies of a morphological, 
genetic, molecular, or of a breeding nature have yet 
been undertaken. 

North American Chara canescens is parthenogenet- 
ic; no males have ever been seen. In Europe and Asia 
there are probably two isolated taxonomic entities with- 
in the broadly designated species, a dioecious male/ 
female sexually reproducing entity of restricted range 
from south-central Europe through central Asia and 
China and the parthenogenetic entity more widespread 
across the continent (Krause 1997). Whether male C. 
canescens can fertilize parthenogenetic C. canescens 
has never been demonstrated. The concept of con- 
specificity is not applicable to parthenogenetic popu- 
lations which is exemplified by a mutant ecorticate 
vegetatively reproducing form of C. canescens from 
Svalbard coexisting in the same springpool as a more 
normal parthenogenetic form (Langangen 2000). 


Vol. 119 


In Newfoundland and elsewhere in North America 
C. canescens and C. evo/uta sometimes inhabit the 
same pond. Although the theoretical potential for C. 
evoluta to fertilize C. canescens may be suggested, 
the morphological differences as described previous- 
ly in this paper remain distinct and no indication of 
intergradation of features has ever been noted in these 
situations. This is strong evidence that the two are 
reproductively isolated and that hybridization does 
not and cannot occur. 


Distribution and Ecology 

Chara evoluta in North America has essentially a 
western distribution, west of the interstate borders con- 
necting the eastern state line of North Dakota with 
the eastern state line of Texas (Figure 4). Almost all 
states west of this line have reports in the literature 
and/or specimens in major herbaria. It is also known 
from southern British Columbia, Alberta, and Saskat- 
chewan in Canada, but not east of Saskatchewan. It 
then reappears in a single disjunct location in coastal 
Rhode Island (Wood 1965; Wood and Palmatier 1954) 
and again in southwestern Newfoundland. Although 
thorough charophyte surveys have not been conducted 
in many areas of North America, some eastern juris- 
dictions have been variously investigated and C. evo- 
/uta has not been recorded, further supporting its 
general absence in the east. These studies include lowa 
(Crum 1975), Illinois (Ebinger and Vogel 1977), Indi- 
ana (Daily 1953), Minnesota, Wisconsin (Prescott 
1962), New York (Wood and Muenscher 1956), West 
Virginia, Virginia, Tennesse, North Carolina, South 
Carolina, Georgia, Alabama, Mississippi, Louisianna, 
Florida (Choudhary and Wood 1973), and others. 

Most reports indicate that C. evo/ura is a species of 
saline coastal or inland waters. Its western distribution 
seems to reflect the climatic conditions of low rainfall 
where rates of evapotranspiration often exceed pre- 
cipitation to produce saline lakes and ponds which are 
often closed systems. In some regions geological de- 
posits are also sources of salts (Hammer 1986). Within 
the stippled region of Figure 4, C. evo/uta only occurs 
locally where such saline conditions exist. East of — 
this region rainfall increases, few inland saline habi- 
tats occur, and all known collections are from coastal 
locations influenced by sea water. 

Inland and coastal saline waters often differ signifi- 
cantly in their ionic composition. For example, Waldsea 
Lake, Saskatchewan, is a magnesium-sodium sulphate 
lake (Hammer 1984) as opposed to marine salinity 
where sodium and chloride predominate as in the 
Newfoundland lagoons. Waldsea water has a conduc- 
tivity and pH in the same range as the Newfoundland 
lagoons and both C. evo/uta and C. canescens are 
known to occur in Waldsea Lake as well as C. Guckel- 
/7iG. O. Allen. Hammer et al. (1983) indicate that the 
Na/Mg ratio plays an important role in governing the 
occurrence of algal species in lakes. C. evo/ufa appears 


2005 


to be capable of adapting to chemically different sati- 
nities, but also to a wide range of salinities. Winter 
and Kirst (1991) and Winter et al. (1996) determined 
that C. canescens has an upper salinity tolerance in the 
mesohaline range of approximately 20 parts per thou- 
sand (ppt) and a lower limit of 1.5 ppt in the low oligo- 
haline range. It may be assumed that this is equally 
probable for C. evo/u/a since both are often associates 
in inland and coastal waters. A few authors report C. 
evoluta from non-saline freshwater habitats (Allen 
1900; Leake 1945). Also V. W. Proctor (personal com- 
munication) indicates that both species can be success- 
fully cultured under low salinity conditions to produce 
viable oospores. Brock (1986), however, suggests that 
for submerged aquatic plants in saline environments, 
extremes of tolerance are not the primary factors in 
determining the flora, but rather evolved life cycle 
mechanisms which allow survival under widely fluc- 
tuating conditions. Thus fluctuating salinities and/or 
ephemeral habitats would foster different floras than 
permanent water bodies with little salinity fluctuation. 

Some degree of salinity and/or particular ionic com- 
position appears to be required by C. evo/ufa (and C. 
canescens), but within this broad tolerance range from 
lower oligohaline (oligosaline) to mesohaline (meso- 
saline), presence or absence in water bodies may be a 
function of biotic community structure. This concept 
suggests that salinity may set the extreme boundaries 
for such species, but that community structure in- 
cluding competition, predation, herbivory, allelopathy, 
and others may determine the colonization ability of 
any given species. Barnes (1999) states that there is 
evidence that brackish water communities are not phys- 
ically structured by salinity, but by biotic processes 
such as predation and competition. Although it is tempt- 
ing to suggest that C. evo/ufa’s apparent absence from 
continental eastern North America is due to the ab- 
sence of any significant saline waters, this may be an 
oversimplification. Even when dispersed, ability to 
establish, grow, and reproduce in complexly evolved 
communities may not be possible despite favourable 
physical parameters. For example, a relationship 
between the presence of certain invertebrate herbivores 
and the distribution of charophyte species has been 
demonstrated by Proctor (1990, 1999). 

In Newfoundland C. evo/u/a grows in association 
with Chara aspera in the five southernmost lagoons 
and Codroy Estuary Pond (Table 2). In these lagoons 
C. aspera is very common, as is C. evo/ufa, but no other 
charophytes occur. This C. evolutalC. aspera associ- 
ation is also recorded from interior British Columbia 
(Allen 1951), Rhode Island (Wood and Palmatier 
1954), and Nebraska (Daily and Kiener 1956). In the 
two northern lagoons (Gravels Pond and St. George’s 
Pond) C. evoluta grows with C. canescens, but C. 
aspera is absent. The C. evolutal C. canescens associ- 
ation is reported from Waldsea Lake, Saskatchewan, 
and Roses Pond (Sweets Lake), British Columbia, 
where they both grow with Chara buckellii, but C. 


MANN AND NAMBUDIRI: CHAROPHYTES OF INSULAR NEWFOUNDLAND II 33 


FiGurE 4. Distribution of Chara evolufa in North America. 
The stippled area indicates its western distribution. 
The two dots indicate the only known eastern sites. 


aspera \s apparently absent. The C. canscens!C. aspera 
combination is recorded for Rhode Island (Wood 1950) 
and reported as a common associate in both North 
America and Europe (Allen 1882; Krause 1997). C. 
canescens has not yet been collected in association 
with C. aspera in Newfoundland, and neither here, nor 
in any other reports have all three species ever been 
reported from the same body of water. Based on two 
well-known ecological principles of competitive exclu- 
sion and of coexistence, it is tempting to try and ascribe 
some ecological significance to this apparent pattern 
of species associations, but such an attempt may be 
premature without a thorough comparison of site con- 
ditions, both biotic and abiotic. 

Chara aspera has a broad North American range, 
commonly occurring in permanent sites from coast to 
coast in southern Canada and tapering to and becom- 
ing more infrequent in Mexico (Croy 1982). It occurs 
more commonly in freshwater situations than either 
C. canescens and C. evoluta, but has an extremely 
broad salt tolerance (Langangen 1974). It is common 
in the Newfoundland upper estuaries of the Little 
Codroy and the Codroy River, however has not been 
collected further north. Ripe oospores have not yet been 
observed for this species in Newfoundland despite 
collections ranging over many years and all summer 
seasons. Dispersal by waterfowl via the many round 
white bulbils produced on the rhizoids of this species 


34 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE 2. Charophytes present in the coastal lagoons/ponds of southwestern Newfoundland investigated in this study. 


C. evoluta 


xX 
xX 


Gravels Pond 

St. George’s Pond 
Codroy Estuary Pond 
Osmond Pond 

Rocky Barachois 
Saltwater Pond 
Second Pond 

First Pond 


KK KK mK MK 


may be considerably more haphazard and infrequent 
than dispersal by spores and may account for its ab- 
sence in the two northern lagoons. However, other 
abiotic/biotic differences as yet undetermined may 
also be involved. 

Chara globularis Thuill. (= C. delicatula Agardh) 
grows in St. George’s Lagoon, the site with the lowest 
recorded salinity. Although this species is normally 
associated with fresh water, it has quite a broad toler- 
ance range and is occasionally reported from low 
salinity waters (Winter and Kirst 1991). It is the most 
common Cara in Newfoundland. 

Relatively few vascular taxa inhabit brackish waters. 
Those associated with charophytes in all the New- 
foundland lagoons include Sago Pondweed, Horned 
Pondweed, and Widgeon-grass. These cosmopolitan 
vasculars of saline coastal and inland waters all belong 
to closely related families and are important food 
sources for waterfowl, as are charophytes. These vas- 
culars are common throughout their ranges wherever 
brackish conditions occur (Melack 1988) although all 
can also occur in alkaline freshwaters (Hammer and 
Heseltine 1988). 

It is well accepted that charophyte spores are readily 
dispersed by waterfowl and shore birds, even great 
distances (Kristiansen 1996). The disjunct Rhode Island 
C. evoluta site may be attributed to such long distance 
dispersal as considerable east-west exchange also 
occurs in addition to the normal north-south migrato- 
ry routes of waterfowl (Bellrose 1976). Undoubtedly 
more coastal populations of C. evo/u/a occur along the 
Atlantic seaboard wherever suitable conditions exist. 
The Newfoundland populations are almost certainly 
a northward dispersal since the last glaciation via the 
Atlantic Flyway. 

North American parthenogenetic C. canescens ex- 
hibits a similar range to C. evo/ura, but tends to have 
a more sporadic occurrence as one moves south from 
western Canada. It ranges up the eastern seaboard into 
Newfoundland, Greenland, Svalbard, and into Europe 
and Asia where many forms and varieties have been 
described. It too is reported occasionally from fresh or 
oligohaline waters (Olsen 1944; Prescott 1962; Winter 
and Kirst 1991; Langangen 2000). Langangen (2000) 
suggests that the reason that C. canescens is not nor- 


C. canescens 


C. aspera C. globularis 


xX 


KX XS XX 


mally found at low salinities in Europe may be due to 
competition from other plants which flourish at low 
salinities, but are excluded at higher salinities. It might 
also be suggested that at the lower salinities, increased 
herbivore pressure may also become a factor (Proctor 
1999). 

As there are no distinct morphological differences 
between C. canescens of the eastern seaboard and that 
of Newfoundland lagoons (Table 1) it can be assumed 
that this species, like C. evo/ufa, entered Newfoundland 
from the south via the Atlantic Flyway. However, be- 
cause of its northern distribution an arctic route can- 
not be ruled out (Mann 1994a). 


Key to Newfoundland Species 

The following key is provided specifically to iden- 
tify the taxa known to occur in Newfoundland and 
Labrador. For an explanation of charophyte structure 
and terminology we recommend Groves and Bullock- 
Webster (1920, 1924), G. O. Allen (1950), and Moore 
(1986). Recommended more general keys to the entire 
North American charophyte flora include Robinson 
(1906) for the Genus Chara, Wood (1948) for the 
Genus re//a, and G. O. Allen (1954) for Genera 
Nitella and Tolypella. Wood (1965, 1967) and Wood 
and Imahori (1964) are essential references for ad- 
vanced studies, but the combining of some species in 
these works discourages the reporting of many good 
taxa. It is recommended that Newfoundland workers 
identify taxa using the currently supplied key and when 
uncertain, consult a specialist. There is always the pos- 
sibility that species new to the province may be dis- 
covered in the future, but such will almost certainly 
be rare or uncommon in our flora. 
la Coronula of 5 cells in one tier, cortical cells 

usually present on the main axis, stipulodes 


present = (Chara) > 3 a.k 6) ons =e 5 
1b Coronula of 10 cells in two tiers of 5, axes 
always without cortication, stipulodes absent ...... 2 


2a Oogonia and oospores round in cross section, 
branchlets consisting of a main axis with 
smaller laterals — 7o/ypella 
Sa ces ee Tee Tolypella glomerata (Desv.) Leonh. 
This is our only known 7o/ypella 
2b Oogonia and oospores somewhat flattened in 
cross-section, branchlets forking usually 
equally one or more times (A722)... haouts ora she titans 3 


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2005 


3a Unbranched end segments of branchlets 
(dactyls) of more than one cell, end cell 
tiny and pointed 
5h tee ee Nitella confervacea (Breb.) A. Braun 
3b Unbranched end segments of branchlets 
(dactyls) of one cell (WV. flevi/rs group) 
4a Plants monoecious (antheridia and oogonia 
ensame plants) 2... /25.%. Nitella flexilis (L.) Agardh 
4b Plants dioecious (antheridia and oogonia 
easepartate plants)... 222. 61...200 Nitella opaca Agardh 
5a Axial cortex composed only of primary rows of 
cells (all rows contain spine cells = haplostichous), 
Axes bristly with many long spines ............... 6 
5b Axial cortex of primary rows with spine cells 
alternating with one or two secondary rows 
without spine cells, Axes not greatly spiny ......... q 
6a Plants monoecious (antheridia and oogonia on same 
plants), coronula width over 180 um 
1 ye Se Ee ee ee Chara evoluta T. F. Allen 
6b Plants only with oogonia, no males with 
antheridia known in North America, coronula 
less than 180 um wide 
Roione. 's, atct: Chara canescens Desy. & Lois. in Lois. 
7a Axial cortex having each primary row 
alternating with one secondary row 
ETT FOS ETCH TC URSIN git Sg a geet Ee ee Bea 8 
7b Axial cortex having each primary row 
alternating with two secondary rows 
GMS OSMCTIONS) ils. taReeh, Sdedtta bata aa 2 ashen eae eae 9 
8a Primary axial cortical rows of cells more 
prominent than the secondary rows 
Be sb ese ucmantn fee em (obi Chara contrarta A. Brown ex Kiitz 
8b Primary axial cortical rows of cells less 
prominent than the secondary rows .. . .Chara vulgaris L. 
9a Plants dioecious, upper and lower stipulodes 
well developed, rhizoids often with tiny 
spherical white bulbils 
OS A ee ee Chara aspera Deth.ex Willd 
9b Plant monoecious, stipulodes especially 
lower ones rudimentary, globular; rhizoids 
a IRE HOTTA eB tsscpatts os c\opaltel By adi ata en afehe ato 10 
10a Primary cortical rows more prominent than 
secondary rows (tylacanthous), upper row of 
stipulodes more elongate than globular lower 
row (sometimes only slightly) 
OE, Sr eae ee Chara delicatula Agardh non Desv.* 
10b Primary cortical row and secondary rows 
equal in size (isostichous), both upper and 
lower stipulode rows of rudimentary 
globular cells 
ey ee Chara globularis Thuill (= C. fragilis Desv.)* 


* These two taxa may show ecologically induced intergrad- 
ing forms and in some treatments are combined under C. 
&lobularis Thuill. They should, however, always be distin- 
guished separately whenever possible until their taxonomy 
can be more precisely defined. 


Discussion 

Insular Newfoundland has a depauperate vascular 
flora compared to the adjacent mainland largely due 
to dispersal difficulty across the straits and to climatic 
and edaphic factors resulting in less diverse habitats 
(Damman 1965). On the other hand, the charophyte 


MANN AND NAMBUDIRI: CHAROPHYTES OF INSULAR NEWFOUNDLAND II 35 


flora compares favorably numerically with many other 
parts of northern North America (Mann et al. 1999). 
It is well known that charophytes are rapid early col- 
onizers of disturbed habitats (Olsen 1944) suggesting 
that dispersal is not a significant limiting factor for 
this group, especially within a single land mass. Almost 
invariably if suitable habitat exists, it will become 
colonized by charophyte species normally occupying 
those particular biotic and abiotic parameters. 

Those Newfoundland taxa with broad ecological tol- 
erances commonly occur across North America, includ- 
ing C. globularis, C. delicatula, C. contraria, C. vul- 
garts, NV. flexilis, and NV. opaca. These are species of 
permanently inundated and relatively stable habitats 
after formation. C. globularis and C. delicatula as 
defined in this paper, in addition to V. /lexi/is and W. 
opaca, are most tolerant of oligotrophic acid waters 
and are ubiquitous across the Island (Mann 1989). Of 
this group, C. de/icat/a is the most tolerant of saline 
conditions and can commonly be found in coastal areas 
subject to some marine influence. C. contraria and 
C. vulgaris are species of high pH waters, commonly 
the limestone areas of the west coast, but also in very 
high pH serpentine pools and of coastal low-saline 
habitats. 

The other portion of the Newfoundland charophyte 
flora is made up of species with more narrowly defined 
habitats, where habitats are rare on the Island and there- 
fore the species themselves are rare. The three species 
featured in this paper (C. evoluta, C. canescens, C. 
aspera) are restricted to soft-bottomed, fluctuating 
saline habitats which are only relatively common on 
the southwest coast. 7o/ypella glomerata 1s restricted 
to coastal freshwater ephemeral sites or their equiva- 
lents which are decidedly uncommon here. W7e//a 
confervacea \s restricted to two west coast sites whose 
common ecological features are still unclear. In all of 
these five rare species, fluctuating and regular habitat 
disturbance are common factors, and all are of coastal 
distribution because this is the only area where habitat 
suitable for their growth occurs. These five rare species 
exhibit a disjunct distribution with the New England 
states. None have yet been reported for the maritime 
provinces or adjacent Quebec. If suitable habitat for 
these species exists in the Maritimes, this disjunct dis- 
tribution may prove to be an illusion of insufficient 
field observation. 

It has been determined that disjunct Newfoundland 
Chara evoluta and Chara canescens populations are 
morphologically and ecologically similar to their coun- 
terparts elsewhere in North America suggesting rela- 
tively recent introductions from the south. The two taxa 
can readily be distinguished by their monoecious/dio- 
ecious (parthenogenetic) conditions respectively and 
by oospore and oogonial features. However, other sub- 
tle differences also may suggest the possibility that at 
least some morphology may be due to convergent adap- 
tation rather than divergent evolution from recent com- 


36 THE CANADIAN FIELD-NATURALIST 


mon ancestors. Their distribution in North America is 
ascribed to their affinity for saline waters. Although 
ecologically similar, there is some indication that their 
niches do not totally overlap and that their salinity tol- 
erance ranges far exceed their actual ecological ranges 
leading one to suspect that biotic factors may be equal- 
ly important in determining presence or absence in a 
given body of water, and therefore their total bio- 
geography. 

The coastal lagoons of south-west Newfoundland 
are unique habitats adding to the provincial biodiver- 
sity by harbouring highly disjunct charophyte species 
within the North American continental perspective. 
Three species, C. evoluta, C. canescens, and C. aspera 
are restricted to this region and have been accorded 
rare status in Newfoundland (Mann 2000*). Along with 
estuaries in the region (Gillespie et al. 1991), lagoons 
provide important feeding and migratory habitat for 
waterfowl and shorebirds, including the endangered 
Piping Plover which nests on the sandy outer beaches 
(Hancock, J. 2001*; Knox et al. 1994). The Banded 
Killifish listed by the Committee on the Status of En- 
dangered Wildlife in Canada (COSEWIC 2000*) as a 
species of special concern in Newfoundland also occurs 
in the lagoons. Much is still unknown about the de- 
tailed flora/fauna and the community structure in these 
saline water bodies. Being sites that can easily be 
altered and degraded by human activity (Martin et al. 
2002; Barnes 1980), their unique features need to be 
further documented in detail and some degree of pro- 
tection should be considered within the developing 
provincial strategy. 


Acknowledgments 

We thank V. W. Proctor, M. V. S. Raju, and A. Lan- 
gangen for their comments and suggestions. Thanks to 
Tony Dickinson for identification of the Banded Kil- 
lifish and to Claudia Hanel for reporting the Codroy 
Estuary site. Greatfully acknowledged are Ellen Block 
and Barbara Thiers for specimen loans from the New 
York Botanical Garden (NY). Loans from the Field 
Museum of Natural History, Chicago (F) were also 
helpful. 

This study was partially funded from the Principal’s 
Research Fund, Sir Wilfred Grenfell College, and a 
Sabbatical Grant from Memorial University. 


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Mann, H. 2000. General status of charophytes of the island 
of Newfoundland. Report for the Newfoundland Rare Plant 
Project, Endangered Species and Biodiversity Section, 


Vol. 119 


Inland Fish and Wildlife Division, Government of New- 
foundland and Labrador, St. John’s, Newfoundland. 


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Received 23 September 2003 
Accepted 21 February 2005 


The Effect of Human Activity on Ant Species (Hymenoptera: 
Formicidae) Richness at the Mont St. Hilaire Biosphere Reserve, Québec 


JONATHAN Z. SHIK!, ANDRE FRANCOEUR?, and CHRISTOPHER M. BUDDLE? 


'Department of Biology, McGill University, 1205 Dr. Penfield Avenue, Montréal, Québec H3A 1B1 Canada; E-mail: jonathan_ 
shik @ hotmail.com 

Centre de données sur la biodiversité du Québec, Université du Québec a Chicoutimi, 637-108 boulevard Talbot, Chicoutimi, 
Québec G7H 6A4 Canada; E-mail: andre_francoeur@uqac.ca 

3Department of Natural Resource Sciences, McGill University, Macdonald Campus, 21 111 Lakeshore Road, Ste. Anne de Bellevue, 
Québec H9X 3V9 Canada; E-mail: chris.buddle @ megill.ca; (author to whom correspondence should be sent) 


Shik, Jonathan, André Francoeur, and Christopher Buddle. 2005. The effect of human activity on ant species (Hymenoptera: 
Formicidae) richness at the Mont St. Hilaire Biosphere Reserve, Québec. Canadian Field-Naturalist 119(1): 38-42. 


The ant (Hymenoptera: Formicidae) fauna of the Mont St. Hilaire Biosphere Reserve, Québec, was surveyed in 2002 and 
2003. Although overall species richness was high, 10 of 40 total ant species collected were limited to anthropogenically dis- 
turbed habitats within the reserve. While only 2 of these 10 species (7esramorium caespitum (L.) and Lasius niger (L.)) can 
definitively be considered introduced, areas altered by human activity (representing a small fraction of the reserve’s total 
area) possess nearly as many unique species as the reserve’s old-growth forest. Although further research will be necessary 
to determine the consequences of such changes in community structure, this study shows the importance of specifying the 
extent of biodiversity surveys within protected habitats to more accurately monitor the effectiveness of conservation efforts. 


Key Words: ants, Hymenoptera: Formicidae, Mont St. Hilaire, anthropogenic disturbance, old-growth forest, species rich- 


ness, inventory. 


Mont St. Hilaire, protected as an International Bios- 
phere Reserve under the Man and Biosphere program 
of the United Nations, is the largest remaining tract of 
old-growth deciduous forest in the St. Lawrence Valley 
(Arii and Lechowicz 2002). Although dominated by 
Sugar Maple (Acer saccharum) and American Beech 
(Fagus grandifolia), this 10 km? old-growth forest con- 
tains diverse habitats including exposed summits rising 
200-300 m above the surrounding forest, low produc- 
tivity bogs dominated by Hemlock (7suga canadensis), 
and stands of Red Oak (Quercus rubra) and Red and 
White pine (Pimus resinosa and P. strobus). Located 
within a suburban setting approximately 32 km east of 
Montréal, Mont St. Hilaire (45°55'N; 75°13'W) also 
contains areas of anthropogenic disturbance. We sought 
to survey the ant species richness (Hymenoptera: For- 
micidae) at Mont St. Hilaire and determine whether 
human activities within and adjacent to the Biosphere 
Reserve have effects on the fauna. 


Methods 

Between 6 May and 31 July 2002, JZS [Jonathan 
Z. Shik] intensively hand-collected ants (243 samples, 
about 7500 individuals) within the boundaries of Mont 
St. Hilaire. Within a given habitat, individual foragers 
and colonies were collected in both hypogeous (e.g., 
under rocks, in leaf litter, and under decaying wood) 
and epigeous (e.g., on living vegetation, along road- 
sides, and on mosses and boulders) microhabitats. 
Additional colony sampling (56 samples) conducted 


by JZS and AF [André Francoeur] from 7 to 10 July 
2003 verified species distributions and yielded two 
species not recorded in 2002 (Brachymyrmex depilis 
(Emery), and Leprothorax ergatogyneus (Francoeur)). 
Voucher specimens have been deposited in the Lyman 
Entomological Museum (McGill University, Macdon- 
ald Campus, Ste. Anne de Bellevue, Québec). 

Because ant distributions vary along abiotic gradients 
of temperature and moisture that depend on exposure 
to sunlight (Brown 1973), ants were sampled in both 
“vegetated” (plant community covers the ground) and 
“exposed” (unshaded rocky) areas. To assess effects 
of human activity within these two habitat groupings, 
both natural and anthropogenically disturbed (hence- 
forth, “disturbed”) habitat were sampled (Table 1). 
Natural vegetated habitat consisted of old growth for- 
est, while disturbed areas were mid-successional fields, 
isolated patches of secondary forest near boundaries, 
and areas dominated by introduced vegetation, espe- 
cially artificially maintained grass areas near build- 
ings. Natural exposed areas were unshaded boulders 
at summits, and rocky outcrops within the forest 
while disturbed exposed areas were roadsides and the 
unshaded gravel of parking lots and a sandy picnic 
area. 


Results 

The ant survey yielded 40 species spread across 17 
genera, making it one of the most species rich locali- 
ties in the eastern Canadian boreal-broadleaved tran- 


39 


ANT SPECIES RICHNESS AT MONT ST. HILAIRE 


SHIK, FRANCOEUR, AND BIDDLE 


2004 


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sition forests (Table 1). Of this total, 10 species were 
most commonly collected (i.e., occurred >90% of the 
sampling observations) in disturbed habitat (disturbance 
specialists), 14 species could be considered specialists 
in natural habitat (>90% of observations in natural habi- 
tats), and 16 are considered habitat generalists, being 
found frequently in both natural and disturbed habitat 
(Table 1). Five of the 10 disturbance specialists (Zep- 
tothorax ambiguus (Emery), Solenopsis molesta (Say), 
Formica lasioides (Emery), Lasius niger (L.), and ZL. 
pallitarsis (Provancher)) were sampled three or less 
times during the course of the study (Table 1). We 
believe this represents true rarity rather than collection 
bias as most of these species were readily detected, 
being found in epigeous habitat. Only Ponera penn- 
sylvanica (Buckley), S: molesta and F- lasioides, might 
have been overlooked in nearby primary forest due to 
being found only under rocks and under litter at the soil 
surface. 


Discussion 

At first glance, ant species richness at Mont St. 
Hilaire—a forest reserve of limited area—is exception- 
ally high. Surveys of natural and disturbed biotopes in 
the much larger areas of Chateauguay and Huntington 
municipal counties in Québec just south of Montréal 
yielded 48 species (A. Francoeur, unpublished data), 
81% of which exist at Mont St. Hilaire. However, when 
considering that only 30 of Mont St. Hilaire’s 40 species 
were found in natural habitats, diversity “within” the 
interior of the reserve is not higher than within other 
similar forests. For example, Letendre et al. (1971) 
found 29 species (22 of which occur at Mont St. 
Hilaire), in post-fire Paper Birch (Bela papyrifera), 
poplar (7opu/us sp.) and Sugar Maple forest at the Uni- 
versité de Montréal field station (9 km), 80 km north- 
west of Montréal. 

Two of the disturbance specialists are considered 
introduced. The presence of Zaszus niger (L.) represents 
a significant extension to its range as it is known as a 
west coast species (Wilson 1955). While collections 
within the city of Montréal suggest that this species is 
established in the region, additional collecting may help 
determine if Z. zZger is introduced from the west or 
occupies an undocumented larger range. 7etramorium 
caespitum (L.), a globally distributed tramp species 
native to Europe (Bolton 1979), was found only along 
roadsides leading to the park’s main entrance. Because 
the other eight disturbance specialists are native to the 
nearctic region, their historical presence or absence 
at Mont St. Hilaire is unknown. It is apparent, however, 
that within habitat modified by humans these ant species 
presently exhibit increased abundance relative to nat- 
ural areas such that within the protected reserve human 
activity has altered ant community structure. 

Anthropogenically disturbed and fragmented habi- 
tat becomes increasingly vulnerable to the establishment 
of introduced ant species (Tschinkel 1988; Suarez et al. 


SHIK, FRANCOEUR, AND BIDDLE: ANT SPECIES RICHNESS AT MONT ST. HILAIRE 4] 


1998). Human traffic could potentially serve as a vector 
for further ant introductions in the reserve’s disturbed 
areas that represent a small percentage of Mont St. 
Hilaire’s total area, but already house a large number of 
unique species—nearly as many as natural areas. 
Although Québec presently lacks the most notorious 
invasive species that are ravaging the native ant faunas 
of other regions (Francoeur 2000; Holway et al. 2002), 
the introduction of any species should be closely moni- 
tored in reserves that are amongst the last remnants of 
their habitat type. While further research will be neces- 
sary to determine the nature of interactions between 
disturbance and natural specialists, we believe that the 
reserve’s old-growth forest ant fauna should be con- 
sidered as a community marked by scattered disturbance 
subpopulations. More generally, these findings illustrate 
the importance of clearly delineating natural from 
anthropogenically disturbed habitat when conducting 
biodiversity surveys within special reserves such as 
Mont St. Hilaire. 


Acknowledgments 

We thank M. Lechowicz, B. Gilbert, E. Lo, H. 
Varady-Szabo, T. Smith, B. Hamel, the Gault Nature 
Reserve, and the Lyman Entomological Museum of 
McGill University. Comments from anonymous review- 
ers improved this manuscript. Funding was provided in 
part by NSERC (M. Lechowicz, Department Biology, 
McGill University, and C. Buddle). 


Literature Cited 

Arii, K., and M. J. Lechowicz. 2002. The influence of over- 
story trees and abiotic factors on the sapling community in 
an old-growth Fagus-Acer forest. Ecoscience 9: 386-396. 

Bolton, B. 1979. The ant tribe Tetramoriini (Hymenoptera: 
Formicidae). The genus 7éframorium Mayr in the Mala- 
gasy region and in the New World. Bulletin of the British 
Museum of Natural History (Entomology) 38: 129-181. 

Bolton, B. 1995. A new general catalogue of the ants of the 
world. Harvard University Press, Cambridge, Massachu- 
setts, USA. 

Brown, W. L., Jr. 1973. A comparison of the Hylean and 
Congo-West African forest ant faunas. Pages 161-185 im 
Tropical forest ecosystems in Africa and South America: 
a comparative review. (Edited by B. J. Meggers, E. S. Avensu, 
and WD. Duckworth). Smithsonian Institution Press, 
Washington, D.C., USA. 

Creighton, W. S. 1950. The ants of North America. Bulletin 
of the Museum of Comparative Zoology at Harvard College 
104: 1-585. 

Francoeur, A. 2000. Document Faunique, no. 1. Entomofaune 
du Québec: Liste des Espéces de Fourmis (Formicides, 
hymenopteres). Corporation Entomofaune du Québec, 
Chicoutimi. 8 pages. 

Holway, D. A., L. Lach, A. V. Suarez, N. D. Tsutsui, and 
T. J. Case. 2002. The causes and consequences of ant 
invasions. Annual Review of Ecology and Systematics 33: 
181-233. 

Letendre, M., A. Francoeur, R. Beique, and J. G. Pilon. 
1971. Inventaire des fourmis de la station de biologie de Il’ U- 
niversité de Montréal, St. Hippolyte, Québec (Hymenoptera: 
Formicidae). Le Naturaliste Canadien 98: 591-606. 


42 THE CANADIAN FIELD-NATURALIST Vol. 118 


Suarez, A. V., D. T. Bolger, and T. J. Case. 1998. Effects of | Wilson, E.O. 1955. A monographic revision of the ant genus 
fragmentation and invasion on native ant communities in Lasius. Bulletin of the Museum of Comparative Zoology 
coastal southern California. Ecology 79: 2041-2056. at Harvard College 113: 1-201. 

Tschinkel, W. R. 1988. Distribution of the fire ants So/enop- 


sis invicta and S. geminate (Hymenoptera: Formicidae) in 
northern Florida in relation to habitat and disturbance. An- Received 20 November 2003 


nals of the Entomological Society of America 81: 76-81. Accepted 28 January 2005 


Seasonal Diets of Newfoundland Martens, Warfes americana atrata 


JOHN W. Gosse! and BRIAN J. HEARN? 


!Terra Nova National Park, Glovertown, Newfoundland and Labrador AOG 2L0 Canada 
2Natural Resources Canada, Canadian Forest Service, P. O. Box 960, Corner Brook, Newfoundland and Labrador A2H 6J3 
Canada 


Gosse, John W., and Brian J. Hearn. 2005. Seasonal diets of Newfoundland Martens, artes americana atrata. Canadian 
Field-Naturalist 119(1): 43-47. 


We analyzed scats (n = 679) and stomach contents (n = 25) collected from 1980-2003 to assess the relative frequencies of 
food types utilized by Newfoundland Marten (A/artes americana atrafa) during summer and winter. Meadow Voles (A/icrotus 
pennsylvanicus) were the most prevalent food item occurring in 80% and 47.5% of samples from summer and winter, 
respectively. Apart from Snowshoe Hares (Zepus americana), which occurred in 28% of winter samples, all other food types 
occurred in <16% of samples during each season. Diet breadth was widest during winter and may be related to a lower avail- 
ability of Meadow Voles during this time of year. 


Key Words: Newfoundland Marten, Martes americana atrata, diet, food habits, Meadow Voles, Microtus pennsylvanicus, 
SCal. 


The Newfoundland Marten (A/artes americana _ island. This study will provide insight into the feeding 
atrata) is one of only 14 native terrestrial mammals — ecology of Newfoundland Marten and will allow future 
on the island of Newfoundland (Dodds 1983) and comparisons of foraging patterns following expected 
is currently listed by the Committee on the Status — changes in small mammal communities. 
of Endangered Wildlife in Canada as Endangered. 

Though this species has been the focus of considerable Study Area and Methods 
research, basic information on seasonal diet patterns Scats were collected opportunistically during field 
of Newfoundland Marten is lacking. studies between 1995-2003 and were air and/or oven- 

The potential prey base for Newfoundland Marten, dried prior to storage. Most scats (70%) were collected 
a genetically distinct subspecies (Kyle and Strobeck near Little Grand Lake in western Newfoundland 
2003) of the American Marten (A/artes americana), (57°50'00N, 48°38'00W) prior to 1999. The remainder 
is limited. Newfoundland has a reduced diversity of | was collected in Terra Nova National Park in eastern 
small mammal prey species (8 species) compared with = Newfoundland (53°50'00N, 48°30'W). Stomachs from 
neighboring mainland areas such as Labrador (17 carcasses turned in by trappers in western Newfound- 
species) and Nova Scotia (23 species; Tucker 1988). land were refrigerated and washed prior to examination. 
Further, only one arvicoline rodent, the Meadow Vole Prey items were identified based on the coloration and 
(Microtus pennsylvanicus), fully overlaps with, and banding patterns of mammalian guard hairs, and on 
occurs in forested habitats used by Marten (Thompson _ the remains of indigestible material including teeth, 
and Curran 1995; Tucker 1988; Sturtevant and Bis- —_ feathers, seeds and insect exoskeletons; a reference 
sonette 1997). collection and an identification manual (Adorjan and 

Home range size of Marten on the island of New- —-Kolenosky 1969) aided in this process. We pooled 
foundland is exceptionally large (Bateman 1986; B. these data with other published (Bateman 1986) and 
Hearn, unpublished data). Gosse (/7 press Journal of — unpublished (Snyder 1986*; Tucker 1988; Drew 
Mammalology) and may reflect the low diversity and 1994) data on food habits of Newfoundland marten 
abundance of small and medium-sized mammals collected from 1980 to 1991 (Table 1). Samples were 
(Lindstedt et al. 1986; Thompson and Colgan 1987). categorized by season (winter, 1 November to 31 
Southern Red-backed Voles (Clethrionomys gapperi), April; summer, | May to 31 October) whenever pos- 


a major food item of marten across their North Amer- __ sible. Prey items were placed into one of 10 categories; 
ican range (Buskirk and MacDonald 1984; Simonet (1) Meadow Voles, (2) Southern Red-backed Voles, 
al. 1999), were first recorded in western Newfound- (3) Masked Shrews (Sover cinereus), (4) Snowshoe 


land in 1999 and their distribution and abundance is — Hares (Lepus americanus), (5) Red Squirrels (7am- 
expanding (Hearn et al. 2005). Herein we report the = asciurus hudsonicus), (6) unidentified mammals, (7) 
diversity of food types consumed by Newfoundland Moose (A/ces a/ces) and Caribou (Rangifer tarandus) 
Marten over an 18-year period prior to the widespread __ carrion, (8) insects, (9) avian remains, and (10) berries. 
establishment of Southern Red-backed Voles on the Samples collected during earlier studies (Bateman 


43 


44 THE CANADIAN FIELD-NATURALIST 


TABLE |. Data source, collection period, and number of 
samples collected. 


Number of 
Data source samples Sampling period 
M. Bateman (1986) 56 1980-1982 
J. Snyder (1986*) pa 1984-1986 
B. Tucker (1988) 194 1986-1987 
G. Drew (1994) 12 1990-199] 
This study 417 1995-2003 
Total 704 1980-2003 


a = stomach contents from carcasses. 


1986; Snyder 1986*; Tucker 1988; Drew 1994) were 
analyzed by the original investigators using similar 
techniques. Contents are expressed as frequency of 
occurrence rather than volumetric estimates to main- 
tain consistency with earlier reporting. Furthermore, 
volumetric estimates of prey items would provide little 
additional information since most scats were entirely 
composed of one food type. Shannon diversity indices 
(Krebs 1998) were calculated for samples collected 
during summer, winter, and for both seasons combined. 
Samples were not partitioned according to sex since 
the sex of the animal depositing the scat was known 
for <15% of the samples. Moreover, previous studies 
indicate that significant differences in inter-sexual diet 
are not apparent (Nagorsen et al. 1989; Nagorsen et al. 
1991; Thompson and Colgan 1990; Simon et al. 1999). 


Results 

A total of 704 samples (679 scats and 25 stomach 
contents) were collected from 1980-2003 (Table 2). 
Meadow Voles were the most prevalent prey item in 
scats and stomach contents, occurring in 70.5% of 
all samples (pooled over sites, seasons, and years). 
Snowshoe Hares were the second most frequent prey 
item. Other food types consumed by Marten included 
Masked Shrews, Red Squirrels, Moose and Caribou 
carrion, insects, birds, berries, and unidentified mam- 
mals; however, overall frequency of occurrence for each 
of these items was <10%. Southern Red-backed Vole 
remains were found in a single scat collected near Little 
Grand Lake in 1999; this coincides with the first record 
of this species on the island of Newfoundland (Hearn 
et al. 2005). Insect exoskeletons were identified as 
vespid wasps. Most bird remains were from smaller 
passerines, however, Ruffed Grouse (Bonasa umbel- 
/us) and Willow Ptarmigan (Lagopus lagopus) were 
identified from samples collected in southwestern 
Newfoundland during the winters of 1980-1982 
(Bateman 1986). Eggshells were also found in several 
scats. Scats contained a variety of berries including 
Creeping Snowberry (Gau/theria hispidula), Wild 
Sarsparilla (Ava/ia nudicaulis), blueberry (Vaccinium 
spp.), Bunchberry (Cornus canadensis), Common 
Raspberry (Awbus ‘daeus), and Three-Leaved False 
Solomon’s Seal (Slacina trifolia). The Shannon 


Vol. 119 


diversity index for the pooled data was 1.6. Eighty- 
eight percent of scats contained only one food item, 
consequently, the importance of larger prey items such 
as Snowshoe Hare and carrion is likely under-repre- 
sented (Poole and Graf 1996; Cumberland et al. 
2001) since these foods result in the production of fewer 
scats due to a lack of indigestible material per volume 
of food ingested (Zielinski 1986). 

Comparisons of summer and winter diet pattern indi- 
cated that Meadow Voles were the most prevalent food 
item throughout the year, though frequency of occur- 
rence was nearly double during summer (Table 2). 
Berries were the second most common food type found 
in samples collected in the summer. In winter, 28% of 
samples contained Snowshoe Hare remains indicating 
a 10-fold increase in use of this food item compared 
with summer. The frequency of Masked Shrews and 
birds was consistent between seasons though both food 
categories occurred in relatively few samples (Table 2). 
All other food types, except for berries and insects, 
which are highly seasonal food items, had higher per- 
centages of occurrence in winter. This more balanced 
occurrence of food types in winter samples resulted in 
a higher diversity index (H’ = 1.89) in this season versus 
summer (H’ = 1.36). 


Discussion 

Newfoundland Marten consumed essentially all pos- 
sible prey species available to them suggesting a gener- 
alist foraging strategy. These findings are in agreement 
with earlier studies (Buskirk and MacDonald 1984; 
Thompson and Colgan 1990; Martin 1994) that collec- 
tively have reported a highly diverse diet, and oppor- 
tunistic use of locally available prey (Nagorsen et. al. 
1989; Nagorsen et al. 1991; Martin 1994). Meadow 
Voles were the most frequent component of the year- 
round diet of Newfoundland Marten and were particu- 
larly prevalent in samples collected from early spring 
to late autumn. Marten commonly consume Meadow 
Voles in other regions (Buskirk and MacDonald 1984; 
Slough et al. 1989; Martin 1994; Poole and Graf 1996; 
Simon et al. 1999), but the frequency of occurrence 
found in samples from this study is the highest known 
to us. On the island of Newfoundland, Meadow Voles 
occur in coniferous forests in addition to open, grassy 
areas (Cameron 1958; Folinsbee et. al. 1973; Thompson 
and Curran 1995; Sturtevant and Bissonette 1997) 
thus their distribution fully overlaps with habitats used 
by Marten. The importance of Meadow Voles may have 
been more pronounced historically, since many of the 
prey species currently utilized by Marten were inten- 
tionally or accidentally introduced to insular New- 
foundland over the last 150 years [1i.e., Snowshoe 
Hare (Dodds 1960), Moose (Pimlott 1953), Masked 
Shrew (MacLeod 1960), Red Squirrel (Payne 1976), 
Southern Red-backed Vole (Hearn et al. 2005)], Ruffed 
and Spruce Grouse (Hancock 1981*). In contrast to 
this study, Marten on the Queen Charlotte Islands and 


2005 


GOSSE AND HEARN: DIETS OF NEWFOUNDLAND MARTEN 45 


TABLE 2. Frequency of occurrence of food items found in 679 Marten scats and 25 stomachs in Newfoundland in winter, 
summer, and both seasons combined (pooled over study sites and years). 


Food item 


Meadow Vole (AW%crotus pennsylvanicus) 

Southern Red-backed Vole ( Clethrionomys gapper!) 
Masked Shrew (Sorex cinereus) 

Snowshoe Hare (Lepus americanus) 

Red Squirrel (Zamiasciurus hudsonicus) 
Unidentified mammal 

Moose (A/ces a/ces)/Caribou (Rangifer tarandus) carrion 
Insects 

Avian remains 

Berries 

Shannon Diversity Index (H’) 


Summer Winter Seasons combined* 
n= 400 n = 236 n = 704 
80 47.5 70.5 
0.25 0 0.1 
10.5 10.6 9.9 
2.8 28 11.4 
1.0 7.6 3.4 
4.3 10.2 4.3 
pie) 8.9 3.6 
2.0 0.4 1.4 
TZ 10.2 8.8 
15.5 0.8 9.7 
1.36 1.89 1.60 


4 Includes samples where the season of deposition was unknown; %frequency of occurrence values do not sum to 100% 


across food items. 


Vancouver Island had a smaller component of small 
mammals in their diet and primarily foraged on birds, 
carrion, and salmonid fishes (Nagorsen et. al. 1989; 
Nagorsen et al. 1991). This ability to utilize locally 
available food sources reflects the opportunistic nature 
of Marten foraging behavior. 

Snowshoe Hares were consumed throughout the year 
but were more prevalent in samples collected during 
winter. This seasonal difference may reflect a decrease 
in the abundance or availability of Meadow Voles asso- 
ciated with snow conditions and a switch to a more 
available prey type. The importance of Snowshoe Hares 
to the winter diet and overall population performance 
of Marten has been documented for Newfoundland 
marten (Bateman 1986) as well as for other northern 
regions (Cowan and Mackie 1950; Raine 1983; 
Thompson and Colgan 1987; Thompson and Colgan 
1990; Poole and Graf 1996). Furthermore, Cumberland 
et al. (2001) documented that larger prey, including 
Snowshoe Hares (8.0%), grouse (12.2%) and Red 
Squirrels (10.8%), represented only 31% of the diet by 
frequency of occurrence but accounted for approxi- 
mately 95% of the caloric intake by marten in New 
Brunswick. We suspect that the introduced Snowshoe 
Hare (Dodds 1983) is now a critical food resource for 
Newfoundland marten in winter, the most energetically 
stressful period annually (Thompson 1986; Buskirk et 
al. 1988). 

In addition to preying on more Snowshoe Hares dur- 
ing winter, Marten increased consumption rates of car- 
rion, Red Squirrels, birds, unidentified mammals, and 
Masked Shrews. This is consistent with the pattern 
observed by Thompson and Colgan (1990) where a 
wider breadth of diet was noted during winters when 
prey abundance was low. Masked Shrews appeared in 
samples for both seasons at similar frequencies but were 
recorded in only 2 of 22 food-habit studies reviewed by 
Martin (1994). Shrews have low body mass and may 
only be utilized in years of food scarcity. Birds are a 
common food source used by Marten across their range 


(Martin 1994) and the frequency of occurrence reported 
here approximated the mean reported from other stud- 
ies (14.5% frequency of occurrence). Similar to 
Snowshoe Hares, carrion likely provided a signifi- 
cant source of biomass, hence total calories, during 
winter. 

The remains of a Southern Red-backed Vole in a 
scat collected from southwestern Newfoundland in 
1999 coincided with the first record of this species 
for the island (Hearn et al. 2005). Scats were not col- 
lected after 1999 in areas where Marten and Southern 
Red-backed Vole distributions overlap (Hearn et al. 
2005), thus we were unable to assess use of this newly 
available prey item. Red-backed Voles are heavily 
utilized by American Marten elsewhere (Buskirk and 
MacDonald 1984; Slough et al. 1989; Thompson 
and Colgan 1990; Simon et al. 1999), and we antici- 
pate increased utilization of this forest vole as its 
distribution and abundance expands. 

Several researchers have noted fluctuating levels of 
prey and subsequent changes in population perform- 
ance of Marten. Thompson and Colgan (1987) report- 
ed reduced population density, enlarged home ranges, 
and lower reproductive performance of Marten dur- 
ing a synchronous decline in principal prey species 
on their study area in northcentral Ontario. Poole 
and Graf (1986) suggested that the Snowshoe Hare 
population cycle has a significant impact on Marten 
populations in the northern boreal forest, and Marten 
harvests in Canada are historically well synchro- 
nized with Snowshoe Hare numbers (Bulmer 1974; 
Fryxell et al. 1999). Similarly, a decrease in small 
mammal numbers in central Labrador was implicated 
in the reduction in female to juvenile ratios and the 
overall harvest of Marten in the following trapping 
season (Simon et al. 1999). It is unknown whether 
Newfoundland Marten are food limited and addi- 
tional research is required to understand the relation- 
ships between fluctuating prey levels and population 
performance for this Endangered species. 


46 


Acknowledgments 

We thank various professional and technical staff, 
in particular Tyrone Mulrooney, John Neville, Rod Cox, 
Cyril Lundrigan, Bill Curran, and Joe Brazil. This study 
was the result of several research projects investigating 
the ecology of the Newfoundland Marten supported by 
Natural Resources Canada — Canadian Forest Service, 
Terra Nova National Park, the Newfoundland and 
Labrador Inland Fish and Wildlife Division, the New- 
foundland and Labrador Forest Service, the Western 
Newfoundland Model Forest, Corner Brook Pulp and 
Paper Limited, Abitibi Consolidated Limited, and World 
Wildlife Fund Canada. 


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John’s, NF. 23 pages. 

Snyder, J. 1986. Newfoundland pine marten carcass exam- 
ination. Newfoundland and Labrador Wildlife Division, 
St. John’s, Newfoundland. 


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Received 17 February 2004 
Accepted 11 February 2005 


Pollination and Breeding System of Lowbush Blueberries, Vaccinium 
angustifolium Ait. and V. myrtilloides Michx. (Ericacaeae), in the 
Boreal Forest 


MasayukI Usut!, PETER G. KEVAN!, and MARTYN OBBARD? 


‘Department of Environmental Biology, University of Guelph, Guelph, Ontario N1G 2W1 Canada (Corresponding author: 
Kevan) 

Ontario Ministry of Natural Resources, Wildlife Research and Development Office, 300 Water Street, Peterborough, Ontario 
K9J 8M5 Canada 


Usui, Masayuki, Peter G. Kevan, and Martyn Obbard. 2005. Pollination and breeding system of lowbush blueberries, Vaccini- 
um angustifolium Ait. and V. myrtilloides Michx. (Ericacaeae), in the boreal forest. Canadian Field-Naturalist 119(1): 
48-57. 


Breeding systems and pollination requirements of two wild lowbush blueberries, Vaccinium angustifolium and V. myrtilloides, in 
the Canadian boreal forest in the Chapleau Crown Game Preserve, Ontario, were tested. Fruit production, size and seediness 
were significantly higher in samples exposed to natural pollination than in those cross- or self-pollinated by hand. There were 
no significant differences among artificial treatments (variously hand-pollinated and bagged) except when cross-pollination 
(xenogamy) was done by insect pins. In V. angustfolium, the density of flowering varied with forest age (canopy closure). It 
was most in open areas and least in the sites with the most mature forest. Although fruit-set and seediness varied among forest 
habitats of different ages, there were no significant differences between sites in forests of different ages. Thus, pollination seems 
to be similarly effective no matter the age of the forest. In both species, fruit-set in 1992, which had severe June frosts, was 
markedly poorer than that in 1993 when the flowers suffered little frost damage. The combined number of complete and 
incomplete seeds from the fruit among the breeding and pollination systems tested were similar; however, the ratio of complete 
seeds to total seeds was greater from cross-pollinated than from self-pollinated flowers. Our observations indicate that there 
is little natural fruit-set without insect-mediated cross-pollination and that cross-pollination provides much better fruit and 
seed-set than does self-pollination. 


Key Words: boreal forest, lowbush blueberries, Vaccinium angustifolium, V. myrtilloides, breeding system, fruit-set, seed-set, 
bumblebees, pollination, pollinators, Charleau, Ontario. 


Lowbush blueberries ( Vaccinium angustifolium and 
V. myrtilloides) are wide spread throughout the North 
American boreal forest. Their fruit is important for 
wildlife (Usui et al. 1994). They grow best under acidic 
and well-drained soils in open habitats (Eck and Chil- 
ders 1966), although they persist and may thrive in 
clearings in darker, old forests. 

Although it is generally accepted that lowbush blue- 
berries usually depend on insects, mostly bees (Hy- 
menoptera: Apoidea) for pollination (Eck and Childers 
1966; McGregor 1976; Reader 1977; Hall et al. 1979; 
Morrissette et al. 1985; Mohr and Kevan 1987; Aras 
et al. 1996; Stubbs and Drummond 1997, 1999, 2001; 
Javorek et al. 2002), most research on pollination is 
from commercial lowbush blueberry barrens and there 
is little information from unmanaged habitats, com- 
pletely natural stands, and especially the boreal forest 
(Kevan et al. 1993; Mohr and Kevan 1987). 

Even in horticultural studies, the effects of self- and 
cross-pollination have not been thoroughly investi- 
gated. Aalders and Hall (1961), and Wood (1968) noted 
a high incidence of self-sterility in V. angustifolium. 
Hall and Aalders (1961) stated that about 5% of the 
plants they observed in Nova Scotia and New Bruns- 
wick were male-sterile and 45% produced less than a 
complete complement of normal pollen. Hall et al. 


(1966) also noted female-sterility associated with | 
breakdown of the integument and the nucellus of the | 
ovules. There is little published information on V | 
myrtlloides, presumably because it is of little commer- — 


cial importance. It is regarded as an obligate entomo- 


philous outcrosser (Vander Kloet and Hall 1981) and | 
well pollinated by various bees (Morrissette et al. 1985; | 


Aras et al. 1996). Reader (1977) examined the breed- 


ing strategies of V myrtil/loides in natural bogs of — 


southern Ontario and reported that this species was 
self-compatible with 40% of flowers setting seed auto- 


gamously, 60-80% setting seeding set when pollination | 


occurred between flowers of the same plant (geito- 
nogamy), and when pollination occurred between two 
different plants (xenogamy), the percentage of flow- 
ers setting seed was 70%. Thus, it is possible that the 
breeding systems of the species are different in differ- 


ent parts of their ranges, as is known for other species of 


plants (Richards 1997). Nevertheless, in nature, pol- 
lination of most Vaccinium spp. by bees is essential 
because of the urceolate (urn-shaped) and pendulous 


flowers, which require that pollinators hang onto the — 


flowers and enter them from below to obtain nectar or 
pollen, or both. The most effective pollinators (Javorek 
et al. 2002), such as bumblebees (Bombus spp. (Api- 
dea)) and an array of solitary bees (Apoidea) (Finna- 


48 


2005 


more and Neary 1978) vibrate the anthers to extract 
pollen (vibratile or “buzz” pollination) (Buchmann 
1983). 

The aim of our study was to examine the effects of 
cross- and self-pollination on fruit-set in these two 
species of wild lowbush blueberries in the boreal for- 
est and the fruiting success of V angustifolium in 
forests of different ages. 


Materials and Methods 

The research was done as a part of blueberry fruit 
productivity project in the Chapleau Crown Game 
Preserve (CCGP), northern Ontario (48°N, 83°W) from 
1990 to 1993. Of the 31 study sites used (Figure 1), 
the main one was Chaplin Number 4, studied during 
the flowering period (20 May to 5 July) of 1993 (Fig- 
ure 2). Chaplin Number 4 site is a typical blueberry 


> % 3 


4 cin 


we 


as OMNR Camp 4 


eat Racine Lake 


UsuL, KEVAN, AND OBBARD: POLLINATION AND BREEDING SYSTEM OF BLUEBERRIES 49 


habitat with acidic, sandy soil covered by organic mat- 
ter. The area was logged until 1990, burned, and the soil 
prepared for tree planting in 1990. Jack Pine seedlings 
were hand-planted in 1991 (Figure 3). 

Twenty-five plots (Im x Im) for V angustifolium 
and 20 plots (Im x Im) for V. myrtloides were 
selected for experiments of the breeding system early 
in the blooming season in 1993 throughout Chaplin 
Number 4 (Figure 4). The plots were separated by at 
least 15 metres to assure studies on different clones 
(Vander Kloet 1988). 

To elucidate the breeding systems of Vaccinium 
angustifolium and V. myrtilloides naturally occurring 
in the boreal forest, five treatments were set in place 
in each plot (1 m x | m) before blooming began. Sprigs 
of blueberries with flower buds were selected for each 
of the following treatments: a) Free: all flowers were 


FicurE |. Location of Chapleau Crown Game Preserve within the boreal forest of Canada (inset) and of 31 study sites for 
lowbush blueberry in the area of the Ontario Ministry of Natural Resources Camp at Racine Lake. 


50 THE CANADIAN FIELD-NATURALIST 


FiGuRE 2. Map of the Chaplin Number of 4 (Chapleau Crown 
Game Preserve) study site, showing the location of the 
900 m? study grid within the burned area (see also 
Figure 3). 


free to be visited by any pollinators; b) Bagged: flow- 
ers were enclosed in a white, fine mesh, bag to exclude 
pollinators; c) Geitonogamy: bagged (as in b) but all 
fresh flowers were individually pollinated by hand 
with pollen from other flowers of the same plant; d) 
Xenogamy: bagged, but all fresh flowers were polli- 
nated individually by hand with pollen from other 
plants more than 15 metres away. The sprigs in each 
plot were marked by pink flagging tape with identifi- 
cation numbers at the base until the harvesting of fruit 
was complete (1 September). 

Three methods were used to transfer pollen grains 
in Vv. angustifolium. toothpicks, cotton balls (approxi- 
mately 2.0 cm in diameter), and Number 4 insect pins. 
For V. myrtilloides, only cotton balls were used because 
there were too few plants to try all methods. Anthers 
from fresh flower(s) were placed on a clean Petri dish 
and cut with toothpicks or insect pins. One torn anther 
was attached to a toothpick or insect pin and then 
touched to a stigma. Fresh, clean toothpicks or insect 
pins were used for each pollination. Individual corollas 
were touched with a small piece of cotton ball and the 
flower was knocked by fingers ten times onto the piece 
of cotton ball. The cotton ball was found to be the quick- 
est and easiest method to use, and toothpicks the most 
difficult. 

Fruit-set of V. angustifolium under different forest 
conditions (age of forest) was compared from 1991 to 
1993. The soils in all these areas were acidic, sandy, 
and covered with a layer or organic duff. Fruit-set was 
measured in total of 100 evenly spaced plots (100 cm? 
= 10 cm x 10 cm) set along 100 m transects from the 
edge of, but within the forest (1 m? = 100 x 100 cm2), 
and into the forest. The study area contained forests 
of all ages from newly regenerating to mature with 


Vol. 119 


30m 


0 
Legend 
Trench (average depth 30 cm) Tree (White spruce) 


A] Burnt logs Tree (Jack pine) 
Stumps Tree (Willow) 


FiGuRE 3. Sketch map of the 900 m? study grid at Chaplin 
Number of 4 (Chapleau Crown Game Preserve) show- 
ing trenches, burnt logs, stumps and living trees. 


closed canopy. All the flowers produced in each plot 
were counted and the fruits later harvested, counted 
and examined for seediness from the same plots during 
1991-1993. These data were compared among forest 
ages and years. 

During the blooming and fruiting season, some 
marked plants were damaged by Moose (A/ces a/ces) 
and Black Bears (l/rsus americanus). To reduce the 
effects of such damage, fresh ripened fruits from all 
sites, treatments, and years were hand-picked every five 
days. Their diameters were measured and the seeds 
counted. 

Before the hand pollination experiments were star- 
ed, a preliminary study of pollen availability and extrac- 
tion from blueberry anthers was made at the start of 
bloom (20 May to 4 June, 1993) to assure that pollen 
was being produced in sufficient quantity for our exper- 
iments. We found that individual anthers from fresh 
flowers (10 anthers per flower) and large buds contained 
from 0 to over 300 pollen grains, thus a blueberry 
flower may contain up to 3000 pollen grains available 
for removal by pollinators (Table 1). The average 
production of pollen grains was 166.95 + 6.03 SE in 
large buds. For experimental pollinations, only fresh 
flowers and large buds which had had no contact 
with insect visitors were used as sources of pollen. 

The weather from 25 May to 5 July in 1993 (the 
main flowering period) was cool and free from severe 
frosts (only 3 times were freezing temperatures, rang- 
ing -1.0 to -1.5°C, recorded at the weather screen at 
the Chapleau station). From 5 July until early Septem- 
ber (the fruiting season) the weather was relatively mild 


2005  USUL, KEVAN, AND OBBARD: POLLINATION AND BREEDING SYSTEM OF BLUEBERRIES a1 


Eat ii! 
Gd a i a 


Pe al ld a a 


604 eal 607 |608 


eh i i dl el il et ot jl ee et ei et al nl a rd nt ld wed 
bid el inde oe lll ld 

Ea Wal! ad kc to 

ies La ad Lee ld ld a 
li i i Kt Se i ia dn dO i i a li i il 
sii lle ed ial ih Vi ne" i di iid“ a ld 
oh al" lt i i ad ld ne i i 


Hoe Wnt dae 2 wn 
i liad ili ell << cle ld 
a 
alee el lee ke be 
gd co iw el ea nd wet a ld 


Ls ic a ie 
oxo Pr f= rs fs or 
ag ee a oe a id os Ee i a," 

Fes a Fy li a at i a Cle alll oi wa 
il iit! i? eb dd ad” We a ool awn eh ad wld 


Cee i ea a i i iS a Wa 
en Sa, a li de Ui a aid ad 
els a i i a Al Mb ng wc i ak i in 
[Tt BS (ft ha ct a a Ba i ia et a is a a 
fi be pl al "Tg i ali ih ele li i a 


8 


BREE 


527 


ld 


327 


Fi a i i i a 


ee Tig di" et ed il 


FiGurE 4. Locations of the 20 randomized plots used for pollination studies within the 900 m2 study grid (Chapleau Crown 


Game Preserve). 


compared to long-term average (an average monthly 

mean temperature was 17.2°C in July and 17.1°C in 

August) and wet (monthly rainfall was 106 mm in 

July and 54.8 mm in August). However, the weather 

in the spring of 1992 was cold with six occasions of 

below freezing temperatures, as low as -4.0 and -2.5°C, 
_ between 21 May and 22 June. 

Statistical analyses of fruit-sets, seed-set, and fruit 
size were made. The numbers of fruits were compared 
in size (mm in diameter) among treatments on the two 
species of lowbush blueberries, respectively. Data 
were subjected to ANOVA and means separated by 
Tukey’s Studentized Range Test at P= 0.05 level (SAS 
Institute 1985), or subjected to a modified ANOVA for 
data sets with unequal variances (Zar 1998 :187) (Toms 
| and Almond, personal communication, see Acknowl- 
| edgments). The latter approach required application of 
| Bonferroni corrections to the critical probability value 


TABLE |. Number of pollen grains extracted from large buds 
of Vaccinium angustifolium collected from four locations 
in the Chapleau Crown Game Preserve. Pollen grains were 
extracted from one anther tube of each flower. 


Range of number Total Mean + SE 
of approximate observations (Number of 
pollen grains Freq. % pollen grains) 
Large bud (grains/anther) 

7), oa 
10-49 Be SH 
50-99 ly 1S 
100-199 64 28.7 
200-299 82. 36:8 
< 300 4520.2 


Total number of 
flowers and anthers 223 166.95 + 6.03 


52 THE CANADIAN FIELD-NATURALIST Vol. 119 
TABLE 2. The comparison of fruit-set ratio among eleven pollination treatments of V angustifolium. 
Number of Total Total Range of 

Number sprigs with flowers fruits Mean + SE fruit-set 

of sprigs fruits (% among among of per sprig 
Treatment examined fruit-set) sprigs sprigs fruit-set % % 
Free (natural condition) 56 56 (100.0) 5570 2837 50.5+2.6a* 12 to 86 
Bagged (no pollination) 25 2 (8.00) 1692 2 OL e-O.le 0 to 1 
Xenogamy using insect pins 8 8 (100.0) 193 40 213243520) Bite Ss 
Xenogamy using cotton balls 16 12 (7) 453 48 10.5 + 2.8b 0 to 38 
Xenogamy using toothpicks 16 & 60:0) 542 ie) 3.4 [3b 0 to 16 
Geitonogamy using insect pins zt ay eT) 302 47 3:T a l.5b 0 to 11 
Geitonogamy using cotton balls —=15 7 ~~ (46.7) 477 26 2.4 +0.8b 0 to7 
Geitonogamy using toothpicks 16 l (6.3) 48] l 0.2 4.0.25 0 to 3 
Autogamy using insect pins 8 3 5 ASH) MZ 12 5,3 + Z.8b 0 to 18 
Autogamy using cotton balls 16 4 (25.0) 262 4 2.0 BOG 0 to 11 
Autogamy using toothpicks 16 l (6:3) 209 1 0.4 +0.4b 0 to7 


*Values in a column followed by the same letter are not significantly different. 


for rejection of our null hypotheses: for Vv. angusi- 
Jolium p < 0.0026; for Ko myrtilloides p < 0.0071 
(Toms and Almond, personal communication). 


In V angustifolium, the fruits that developed from 
free pollinations had the greatest number of seeds (Table 
4), but those that developed from the hand-done xeno- 


gamy had significantly fewer (ANOVA F, ,, = 7.26, 


Results p = 0.05), and those from hand done geitonogamy and 
In V. angustfolium (Table 2), the fruit-sets, under pol- autogamy were by far the least and were not signifi- 
lination regimes were significantly different (F,,,,= 128, cantly different from each other (ANOVA F, ,, = 1.08, 


P<<0.001). Significantly the least amount of fruit-set 
was obtained from flowers enclosed in bags and there- 
after untouched (2 fruit from 1692 flowers). The most 
fruit was produced by the flowers open to natural pol- 
lination. Although artificial xenogamy produced more 
fruit than did autogamy or geitonogamy there were no 
significant differences in fruit-set among hand-pollinated 
and bagged treatments, regardless of the method used 
(t ranged from 0.6 to 4.8; p ranged from 0.8 to 0.1). 
Our statistical analysis indicates that artificial xeno- 
gamy done with an insect pin was not significantly 
different from the open pollinated controls in terms of 
fruit-set (t = 6.7; p= 0.12). In V myrtlloides (Table 3), 
fruit-sets under pollination regimes were significantly 
different (F,,, = 17; p << 0.001). Again, open polli- 
nation produced more fruit than any of the other four 
treatments among which there were no significant dif- 
ferences (t ranged from 1.1 to 1.9; p ranged from 
0.31 to 0.47) except for bagged and left alone, which 
again had markedly few fruits (6 from 1794 flowers). 


p = 0.05; F, |) = 1.26, p = 0.05). Fruit diameters were 
also affected by treatment (Table 4). Free pollinations 
resulted in the largest and seediest fruits (31.3 + 5.9 
SE seeds in the 12.5 mm diameter class of fruits) with 
the other treatments significantly different from each 
other and the average size of fruit being smaller (Table 
4) (ANOVA Fie = 8.20, p = 0.05). The fruits irom 
geitonogamy and xenogamy were disproportionately 
smaller with fewer seeds than those from free polli- 
nation. There were no significant differences in fruit 
size and seediness from flowers pollinated by geito- 
nogamy and autogamy (Table 4). The few fruits from 
flowers bagged and left along were very small (5 mm 
in diameter) and seedless; they were omitted from 
analyses. 

In comparing seediness and fruit size (in 1 mm diam- 
eter classes from 4.5 to 12.5 mm) of free pollination and 
xenogamy treatments on V angustifolium, some regres- 
sion relationships were calculated and found to be signif- 
icantly different from zero (Table 4 and Figure 5). How- 


TABLE 2. The comparison of fruit-set ratio among five pollination treatments of V angustifolium. 


Number of Total Total Range of 

Number sprigs with flowers fruits Mean+ SE _fruit-set 

of sprigs fruits (% among among of per sprig 
Treatment examined fruit-set) sprigs sprigs fruit-set % % 
Free (natural condition) 38 38 (100.0) 4201 2336 57,85.£03a* Il to 86 
Bagged (no pollination) 18 & yet 22:2) 1794 6 O22 01e" . Oto d 
Xenogamy using cotton balls 18 1] (61.1) 602 61 Ma 4.7b “Oto 73 
Geitonogamy using cotton balls 17 6 4.53.3) 590 he 2.1+0.9bb Oto9 
Autogamy using cotton balls 17 arer(23.5) 209 11 5322.50" “Oto S51 


*Values in a column followed by the same letter are not significantly different. 


2005 


UsuL, KEVAN, AND OBBARD: POLLINATION AND BREEDING SYSTEM OF BLUEBERRIES 53 


TABLE 4. The comparison of the complete seed production and fruit size produced from 4 pollination treatments of V 


angustifolium. 
Fruit Size Free Xenogamy Geitonogamy Autogamy 
Number Mean+SE Number Mean+ SE Number Mean + SE Number Mean+SE 
diameter of of complete of of complete of of complete of of complete 
(mm) fruits seeds fruits seeds fruits seeds fruits seeds 
LS S 31.2502 5.918a" 
Ps 10 ~ 26.300 =:3.173a 
10.5 22 25.000'2.764a 3 16.000 + 8.737a 
9.5 18 =. 22.611 +2.836ab 11 8.455 +2.436ab 4 3.750 + 0.854a 
8.5 24 13.8752 1.989be 26 +.4.192 £0433bc 1 2.000a l 1.000a 
LES LS. “10,200 21-5226 31° 3.39320 H47c 10. ~—-: 1.600 + 0.267a 7 2.286 + 0.680a 
6.5 14 =10.143 + 1.421c 15. 2.033 :/0.236¢ 3 1.333. + 0.333a 2 1.500 + 0.500a 
=) 10 9.500 + 3.027c S 2000 0:378c a 0667 £0:333a 
4.5 6 1.833 + 0.654c 2 2.000+ 1.000a 
Total 121 100 19 14 
F - values and df: E413 = 8.20; F, 9; = 7.26; F, ,,= 108; F, j9= 1.26; ns 
p < 0.05 p < 0.05 p < 0.05 


*Values in columns followed by the same letter are not significantly different 


ever, no regression relationship was found for geito- 
nogamy and autogamy mostly because of the small 
number of fruits (for example, r? = 0.023 and 0.039 for 
linear regression for each treatment respectively). The 
comparison regression lines between free pollination 
and xenogamy were significantly different from each 
other for all models, i.e., the regression line in free polli- 
nation is higher than that in xenogamy. 

Between sites in CCGP, percent fruit-set and seed- 
iness of V angustifolium did not differ significantly 
with different forest ages (Tables 5, 6) even though 
the numbers of flowers and fruits in younger forests 
were greater than those in the older forests (ANOVA 
F, , = 0.16 in 1991, F, ,, = 0.70 in 1992, F,, = 0.32 in 
1993, and F,,, = 0.22 on average for 1991 to 1993, 
inclusive). For example, there were 683 ripened fruits 
from 2108 flowers per | m? during 1991 to 1993 
from patches in the young forests (A: 1 to 5 year-old 
after clear-cut or burned), compared to 63 ripened 
fruits from 190 flowers per 1 m? in old forest sites (E: 
36 to 100 year-old forests after clear-cut), during the 
Same years; these differences were statistically sig- 
nificant (ANOVA F, ,,= 20.67, for flower production, 
and F, ,,= 5.42, for fruit production). 


Discussion 

The urceotate flower structure restricts flower visitors 
and the most effective pollinators are large buzz-pol- 
linating bees such as bumblebees (Bombus spp.) and 
larger andrenids (Avdrena spp.) (Finnamore and Neary 
1978; Mohr and Kevan 1987; Stubbs and Drummond 
1999, 2001; Javorek et al. 2002), although other, commer- 
cially available, bees (e.g., Apis mellifera L. (Aras et al. 
1996; Javorek et al. 2002) and Megachile rotundata 
F. (Stubbs and Drummond 1997)) may be useful. In fact, 
there was almost no fruit-set without manipulation of 
some sort on the flowers of Vaccinium angustifolium 


and V. myrtilloides (Table 2 and 3). Natural pollination 
resulted in 12 to 86% fruit-set in V. angustifolium and 
11 to 86% fruit-set in V myrt/loides. Cross-pollina- 
tion by hand resulted in a fruit-set up to 38% in Vv 
angustifolium and 73% in V. myrtilloides. From selt- 
pollinations, fruit-set was low (less than 18% and 31% 
in both species, respectively). 

Fruit size and seediness were greater in the natural 
conditions of free pollination. In the experiments, the 
few fruits produced by self-pollination were small with 
few seeds (1 to 6); more complete seed production was 
obtained by cross-pollination by hand (1 to 33 seeds), 
but even this was low by comparison by open pollina- 
tion (Table 4). Eck and Childers (1966) also noted that 
natural pollinations produced larger berries and more 
seeds than did artificial cross-pollinations. Blueberry 
stigmata remain receptive for 5 to 8 days (Merrill 1936; 
Moore 1964; Wood 1962), however, fruit-set is not initi- 
ated if pollination does not occur within 3 days of anthe- 
sis (Chandler and Mason 1935). Thus, the longer period 
of pollination and greater amounts of pollen available 
under natural conditions would be expected to give rise 
to more, larger, seedier fruits. 

Our results confirm the importance of natural xeno- 
gamy by insect pollinators in the fruit-set and seed-set 
of V angustifolium and V. myrtitloides in the boreal for- 
est of the CCGP. The small amount fruit and seed-set by 
experimental geitonogamy, autogamy and bagging alone 
may be the result of small amounts of wind-dispersed 
pollen (bagging alone) or accidental contamination or 
both (geitonogamy and autogamy). 

Fruit-set ratio under natural, free pollination, was 
variable, ranging from 11% to 86% in V. angustifolium 
and V. myrtilloides. This suggests that many flowers are 
not pollinated or that other factors operate such as male- 
and female-sterility, or both (Aalders and Hall 1961; 
Hall and Aalders 1961; Hall et al. 1966; Reader 1977). 


54 THE CANADIAN FIELD-NATURALIST Vol. 119 


1) Free pollination 


High 
~ Mean + SE 
Low 


Number of complete seeds 


45 5.5 6.5 7.5 8.5 9.5 10.5 11.5 125 
Fruit size (mm in diameter) 


2) Xenogamy 


High 
+ Mean + SE 
Low 


Number of complete seeds 


4.5 5.5 6.5 7.5 8.5 9.5 10.5 11.5 12.5 
Fruit size (mm in diameter) 


FIGURE 5. Comparison of several regression models of seediness on fruit size between free pollination and xenogamy for 
Vaccinium augustifolium in Chapleau Crown Game Preserve in 1993. 1) Free Pollination: Linear regression: r? = 
0.3061, Y = -13 + 3.6X; Exponential regression: r? = 0.3297, Y = 1.6e°7**; Logarithmic regression: r? = 0.2904, Y = -46 
+ 29.5 (logX); Power regression: r? = 0.3296, Y = 0.15X*!, N = 121. 2) Xenogamy: Linear regression: r* = 0.2156, 
Y = -9.0 + 1.7X; Exponential regression: r? = 0.3436, Y = 0.2e°*>*; Logarithmic regression: r? = 0.1839, Y = -19 + 11.3 
(logX); Power regression: r? = 0.3221, Y = 0.02X?4, N = 100. 


2005  UsuL, KEVAN, AND OBBARD: POLLINATION AND BREEDING SYSTEM OF BLUEBERRIES 55 


TABLE 5. Comparison of flower and fruit production and percent fruit-set among V. angustifo/ium in forests of different ages 
from 1991 to 1993. Forest age groups (years after clear-cut or burned): A (1-5); B (6-10); C (11-20); D (21-35); E (36-100). 


Year and Number of Number of Flowers/m? Number of Fruits/m? % Fruit-set/m? (fruits/flowers) 
forest age Observations SoS aa a lm Sen a 9 RPS See a Pan ARN Dicken Oran rca 
group (Sites) Mean + SE Mean + SE Mean + SE 

1991 . 

A 3 1493.0 + 181.18a 414.67 + 15.84a 28.67+4.4la 

B 3 652.0 + 192.99b 225.00 + 78.36 ab 33.00 + 8.72a 

c 0 — — — 

D 3 104.7 + 17.05b 36.67 + 16.83b A335 +°23:15a 

E | 124.0b 37.00b 30.00a 
F-value and df: F,,=15.57;p<0.05 F,,=12.30;p<0.05 F, ,= 0.16; ns 1992 
1992 

A 4 2169.5 + 535.03a 424.00 + 206.47a 18.00 + 5.79a 

B 2 1721.0 + 113.00ab 352.00 + 145.00a 21.00 + 10.00a 

C 3 962.3 + 236.30ab 247.00 + 85.78a 26.00 + 5.13a 

D 2 140.3 + 20.73b 19.00 + 4.04a 13.67 + 2.73a 

E 4 118.8 + 42.63b 29.50 + 14.80a 21.75 + 4.25a 
F-value and df: F,,, = 8.92; p< 0.05 Bat 2-14 DS F,,, = 9.70; ns 1993 
1993 

A B 2642.7 + 167.41a 1297.33: 157:79a 48.67 + 3.18a 

B 2 1403.5 + 160.50b 590.00 + 286.00ab 44.50 + 25.50a 

C 4 1053.8 + 212.30b 410.00 + 116.66b 38:25:% 7.252 

D ] 415.0b 142.00b 34.00a 
E Z 366.5 + 166.50b 144.50 + 132.50b 29.00 + 23.00a 
F-value and df: F,,= 17.54; p< 0.01 F,,= 7.62; p< 0.05 F,,= 0.32; ns 
Average during 1991 and 1993 

A 10 2108.5 + 254.02a 683.20 + 159.13a 30.40 + 4.95 a 

B 7 1172.1: 208.27b 365.57 + 97.94ab 32.86 + 7.73a 

i 7 1014.6 + 145.59b 340.14 + 77.61ab 33.00 + 4.99a 

D vi 164.3 + 43.52c 44.14 + 17.89b 28.86 + 10.38a 

EB 7 190.3 '62'53¢ 63.43 + 36.58b 2500 23,91 2a) 
F-value and df F 43> 20.67; p < 0.01 Fy 43= 5.422005 Eyes —=i0s2 eons 


*Values in column followed by the same letter are not significantly different (Tukey’s Studentized Range Test at P< 0.05). 


TABLE 6. The comparison of the seediness of naturally pro- 
duced fruits on V. angustifolium at six study sites. 


Study Site Number Number of 

of fruits seeds/fruit 

measured Mean + SE 
Clifton Number | 18 22.667 + 2.894a* 
Clifton Number 2 12 18.500 + 3.115a 
Clifton Number 3 49 19.265 + 2.045a 
Copperfield Number 3 9 13.222 + 3.894a 
Copperfield Number 5 3 13.615 +2.793a 


Copperfield Number 9 20 16.150 + 2.404a 


*values in column followed by the same letter are not signifi- 
cantly different (Tukey’s Studentized Range Test at P< 0.05). 


In 1993, we found a few strange flowers (e.g., no pistil 
(one sample), short style nearly the same length of sta- 
mens (two), three-divided stigmata (one), and little or 
no pollen within the anthers (Table 1)), but the incidence 
of these aberrations is too low to explain all our records 
of diminished fruit and seed-set under natural pollina- 
tion. Nevertheless, the fruit-sets we observed under 
natural conditions (means: 50.5% in V. angustifolium 


and 57.8% in V. myrtilloides) at Chaplin Number 4 
and other study sites in 1993, indicates that the wild bee 
populations (or pollinator forces (Kevan et al. 1986)) 
were ample by comparison with those on some commer- 
cially managed lowbush blueberry barrens (an excellent 
commercial crop for a fruit-set of 50%, but it is usually 
less 10 to 20% (Karmo 1957)). Our results are different 
from those obtained by Reader (1977) for Vo myr- 
tilloides. The importance and variability in fruit-set in 
open pollinated flowers are demonstrated by both 
studies, with similar results. However, Reader’s results 
indicate much greater capacity for self-pollination 
(autogamy and geitonogamy (60—80%)) in the plants 
he studied than we found (O-31%)). That large discrep- 
ancy suggest that the two populations, his in two bogs 
in southern Ontario and ours in the boreal forest, have 
different sexual reproductive strategies. Perhaps the 
relatively isolated and smaller populations in southern 
Ontario have adapted by having lesser reliance on 
entomophily in the face of less diverse guilds and small- 
er populations of pollinating bees; Richards (1997) pro- 
vides examples of other plants with variable sexual 
reproductive strategies. Nevertheless, fruit-set varies 


56 THE CANADIAN FIELD-NATURALIST 


from year to year and location to location. Weather dur- 
ing bloom and the size of the pollinator force may be 
important. At CCGP, 1991 and 1993 were relatively 
good for blueberry production because there was little 
severe frosty weather to damage the flowers in June 
(Environment Canada 1994). However, in 1992 there 
were several severe June frosts that killed many flowers 
and buds, especially in the Clifton Number | study site 
where from 1800 flowers only 12 fruits resulted vs. 1094 
fruits from 2540 flowers in 1993. At Clifton Number 
1 the young open, forest is susceptible to severe June 
frosts by advective cooling (Cochran 1969a). 

Our hypothesis that forest age affects fruit-set and 
seed-set was not validated. We found no significant dif- 
ferences in percent fruit-set among the forests at differ- 
ent ages, but did note significantly fewer flowers per unit 
area in older forests than in open habitats (Table 6). 
Pollinator populations were not as great in older forests 
sites as in the opened area (Usui 1994), however, we 
observed some shade-loving bumblebees such as &. 
vagans in the former (Usui 1994) as also noted by 
Heinrich (1979) in Maine. The relative paucity of flow- 
ers and paucity of pollinators seem to offset each other 
to allow for adequate pollination and pollinator forage. 
As suggested in Table 6, some locations, which have 
relatively larger fruits and more complete seed produc- 
tions, have greater bee diversity and population. 

During the blueberry flowering season at CCGP 
and Chaplin Number 4 study site during 1993, many 
bees (53 species (Usui 1994)), especially the bumble- 
bees, Bombus ternarius and B. rerricota, visited many 
flowers in rapid succession. They showed several 
flower-searching patterns; e.g., mainly toward the sun 
or windward in straight lines, in zig-zags, or in circles. 
For example, one queen of B. sevnarius visited 1297 
flowers among 55 V. myrtilloides plants within 30 m? 
(5 m X 6 m) in only 24.5 minutes (1600 h on 16 June) 
flying roughly in two large circles and re-visiting the 
same inflorescence of the plants (Usui 1994). Such 
behaviour would promote cross-pollination among 
different clonal plants as she transfered many differ- 
ent sources of viable pollen grains in a short time. 


Acknowledgments 

We appreciate the assistance of George Kolenosky 
of Ontario Ministry of Natural Resources for offering 
an excellent study opportunity during 1990 to 1993 in 
the Chapleau Crown Game Preserve. We also appreciate 
the assistance given by Diana Philbrick, Department 
of Nutritional Sciences, University of Guelph. We are 
especially grateful to Judith Toms and Steve Almond, 
University of Waterloo for statistical assistance and 
advice. 

Financial support from NSERC (Operating/Discov- 
ery Grants to P. G. Kevan) and from OMNR Grants 
and funding to P. G. Kevan and M. Obbard. 


Vol. 119 


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Received 13 February 2004 
Accepted 27 January 2005 


Productivity of Osprey, Pandion halraetus, Nesting on Natural and 
Artificial Structures in the Kawartha Lakes, Ontario, 1991-2001 


PAMELA A. MARTIN!, SHANE R. DE SOLLA!, PETER J. EWINS*, and MICHAEL E. BARKER? 


'Canadian Wildlife Service, Environment Canada, Box 5050, 867 Lakeshore Road, Burlington, Ontario L7R 4A6 Canada 
*World Wildlife Fund, 245 Eglington Avenue East, Toronto, Ontario M4P 3J1 Canada 
3Wetland Habitat Fund, 1289 Algonquin Blvd., Peterborough, Ontario K9H 6N1 Canada 


Martin, Pamela A., Shane E. de Solla, Peter J. Ewins, Michael L. Barker. 2005. Productivity of Osprey, Pandion haliaetus, 
nesting on natural and artifical structures in the Kawartha Lakes, Ontario, 1991-2001. Canadian Field-Naturalist 
119(1): 58-63. 


Ospreys (Fandion haliaetus) declined throughout the Great Lakes basin during the 1950s to 1970s due to usage of organochlo- 
rine pesticides. Following the banning of DDT in 1972, artificial elevated nest structures were erected in the Kawartha 
Lakes region of south-central Ontario to aid in their recovery. As the population grew, large stumps of flooded trees, < 1 m 
above the surface of the water became important nesting sites, despite their propensity to flood in turbulent weather conditions. 
We compared the productivity of Ospreys among nest substrates and longevity of the nests in this area from 1991 to 2001. 
Of 260 individual nesting attempts made over the 11 years, 57% used man-made structures, primarily either quadrupod 
nesting platforms or utility poles. Of nests on natural substrates, stump nests accounted for 37% of total nesting attempts; 
elevated tree nests were relatively uncommon (6%). Productivity of stump nests was significantly greater than that of artificial 
or tree nests (1.48 versus 1.16 and 0.73 chicks produced per occupied nest, respectively). Nevertheless, survivorship of stump 
nests was less than that of platform nests after 3 years of age, as high water levels, storms or winter ice activity destroyed some 
of these low nests between breeding seasons. Ospreys were able to attain greater productivity in these stump nests than on 


man-made nesting substrates. 


Key Words: Osprey, Pandion haliaetus, nest success, productivity, nest platforms, Ontario. 


Osprey (Handion haliaetus) populations in the Cana- 
dian Great Lakes basin underwent a severe decline in 
response to organochlorine pesticides between 1950 
and 1972; however, numbers had already been deplet- 
ed earlier in the century due to habitat loss through 
deforestation and the onset of intensive agriculture 
and urbanization (Ewins et al., 1995). As populations 
gradually recovered in the 1970s following the ban on 
the use of DDT, Ospreys began using a wide variety of 
man-made structures, such as utility poles, television 
antennae and navigation markers, as support for their 
bulky stick nests (Ewins 1996). In addition, Ospreys 
readily utilize custom made nesting platforms, typi- 
cally located on quadrupods or tripods, 3-5 m in height, 
embedded in the sediment in the littoral zones of lakes, 
or mounted on disused utility poles on rocky islands 
(Ewins 1996). By the mid 1990s, 36% of occupied 
Osprey nests in the Great Lakes basin were on artifi- 
cial structures. 

In 1978, the Ontario Ministry of Natural Resources 
began monitoring a recovering Osprey population in 
the Kawartha Lakes region of south-central Ontario. 
During the course of the study, artificial nesting plat- 
forms were erected for their use. In 1991, the Cana- 
dian Wildlife Service initiated a study to determine if 
Ospreys breeding on the Great Lakes were continuing 
to suffer any adverse effects of contaminants, and in- 
cluded several lakes within the Kawartha Lakes system 
to provide comparative data on what was assumed to 


be an uncontaminated inland breeding population 
(Ewins et al. 1995; Martin et al. 2003). A colony of 
Ospreys breeding at high density in Sturgeon Lake was 
unexpectedly found to be more highly contaminated 
with PCBs than birds in other areas and monitoring 
there continued until 2001 (Martin et al. 2003), pro- 
viding 11 seasons of data on reproduction and nest 
success relative to various nest substrates. 

Nests in the northwestern Kawartha Lakes, includ- 
ing Sturgeon Lake, were primarily located either on 
artificial nesting platforms or on large flooded stumps 
which protruded less than 1 m above the surface of 
the water. A minority of nests was located in trees or 
on utility poles. Artificial nest sites of Ospreys in the 
Great Lakes basin have been shown to have slightly 
higher productivity and lower collapsing rates com- 
pared to natural nest sites (Ewins 1996). Similarly, in 
Saskatchewan, rates of nest success were greater on 
artificial structures than natural structures, though pro- 
ductivity was only marginally greater (Houston and 
Scott 1992). Productivity of Ospreys nesting on arti- 
ficial sites in Baja Peninsula, Mexico, was higher than 
those that used natural nest structures (Castellanos 
and Ortega-Rubio 1995). Thus, we expected that pro- 
ductivity and success would be lower in nests on natural 
compared to artificial structures. Also, natural nests 
may be more transitory than artificial nests, so we con- 
trasted longevity of nest use between natural and arti- 
ficial nests. 


58 


59 


2005 MARTIN, DE SOLLA, EWINS, AND BARKER: OSPREY NESTING 


Lake Dalrymple 


Balsam Lake 


Lovesick 
Lake 


Mitchell Lake 
Cameron Lake 


Canal Lake 


Buckhorn 
Lake 


Emily 
Sturgeon Lake Creek 
SS 


colony 


Study Area 


Sturgeon 
Lake 


Pigeon 


ate Chemung 


Lindsay 


Lake Scugog 


Lake 


(@) Peterborough 


FiGuRE |. Kawartha Lakes study area indicating its position in the Great Lakes basin and the Sturgeon Lake Osprey breeding 


colony. 


Methods and Materials 

The study area was a subset of lakes within the 
Kawartha Lakes region in southcentral Ontario, a 
transitional zone bordering the Canadian Shield and 
the Great Lakes-St. Lawrence Lowlands ecozones. 
The Kawartha Lakes are a chain of 17 shallow, main- 
ly eutrophic lakes interconnected by a series of rivers 
and man-made canals. The water levels are controlled 
in some areas through the use of dams, and the littoral 
regions of some lakes were flooded as a result of the 
construction of the Trent-Severn Waterway over the 
last two centuries 

A group of Kawartha Lakes at the northwest end of 
the chain was surveyed from 1991 to 1996; these in- 
cluded Balsam, Canal, Dalrymple, and Mitchell Lakes 


and Sturgeon Lake, which included two main nesting 


_ areas: Emily Creek and a bay in the south end (Figure 


1). The south end of Sturgeon Lake was the site of a 


_ fairly dense nesting “colony” of Ospreys, at which as 


many as 17 occupied nests occurred within a 3.5 km 
radius. Monitoring of the Sturgeon Lake colony con- 
tinued until 2001. Because surveys at Sturgeon Lake 
continued an additional 5 years (1997 to 2001) beyond 
those of the other Kawartha Lakes (1991-1996), we 
tested to determine if productivity was similar before 
and after 1997 using a two-factor ANOVA, with nest 


substrate and time period (before 1997 and after 1997) 
as factors. There were no differences in productivity in 
relation to either factor or their interaction (p > 0.05), 
so further analyses included data from Sturgeon Lake 
for all years. 

Aside from three nests in dead trees, natural nests 
all consisted of stumps of trees that had been felled 
prior to the flooding of land surrounding the original 
lake basin. One stump nest was later modified to in- 
clude a box to improve stability, but was nevertheless 
considered to be a natural nest. The remaining nests 
were artificial and consisted of utility poles (includ- 
ing one unused television antenna), platforms raised 
on quadrupod support poles, or duckblinds. Nests 
were checked twice a year by boat. The first visit was 
in late May during incubation to count clutch size and 
determine occupancy. Nests on elevated man-made 
structures were accessed using an aluminum exten- 
sion ladder. The second visit was in late June or early 
July to determine productivity and band chicks. Nests 
were Classified as successful if at least one chick sur- 
vived to 5 weeks old. Productivity was calculated as 
mean number of chicks fledged per occupied nest. 
Productivity and longevity of nests were compared 
among nest substrates using analysis of variance 
(ANOVA). 


60 THE CANADIAN FIELD-NATURALIST 


Frequency tables were used to compare productivity 
and occupancy rates, both among years and among 
nest substrates. Pearson x7 was used to test for differ- 
ences in frequency. Survival analysis was used to deter- 
mine if the longevity of nests, defined as the number 
of years that a nest was occupied, differed among nest 
substrates. For the survival analysis, the first year that 
a nest was occupied was treated as year 0. The Kaplan- 
Meier method (Kaplan and Meier 1958) was used to 
estimate the survival function, and Gehan’s general- 
ized Wilcoxon test and Cox’s F test were use to con- 
trast survival times among nest types. Statistica 5.5 
was used for statistical analyses (StatSoft Incorporat- 
ed 2000). Statistical significance was inferred at the 
P < 0.05 level. 


Results 

During 11 years of monitoring a northwest portion 
of the Kawartha Lakes region, 260 individual Osprey 
nesting attempts were made (Table 1). Of these, 43% 
were on natural substrates and 57% were on artificial 
structures. Of the natural substrates, overwater flooded 
stumps accounted for the majority, and were used in 
37% of all nests; in contrast, trees provided substrate 
for only 6% of all nests (Table 1). The most commonly 
used artificial structures were the quadrupod nesting 
platforms (33% of all nests), followed by utility poles 
(20%) and duck blinds (4%). 

There were no differences in productivity among 
the 5 nest substrates overall (Tree, Stump, Platform, 
Utility Pole, Duckblind: P = 0.09). Nevertheless, dif- 
ferences were found when nest substrates were com- 
pared within artificial and natural categories. Produc- 
tivity was higher in Stump nests compared to Tree 
nests (P = 0.05); however, there were no differences 
among artificial nest substrates (P = 0.51). Substrates 
were then pooled into three groups: Stump nests, Tree 
nests, and Artificial nests. Productivity varied among 
these three nest substrates (P = 0.04, Figure 2), and 
was higher in Stump nests than either Artificial or 
Tree nests, having 1.48, 1.16 and 0.73 fledged chicks 
per occupied nest, respectively. 

To test effects of nest height on productivity and 
nest success, we grouped nests according to height: 


Vol. 119 


Productivity 


Stump Tree 


Artificial 


FIGURE 2. Productivity (number of chicks per active nest) of 
Artificial, Stump and Tree nests of Ospreys in the 
Kawartha Lakes region, 1991-2001. 


those situated greater than 2 m above water level were 
High nests (Platform, Tree, and Utility pole nests), and 
those less than 2 m above water level as Low nests 
(Stump and Duck blind nests). As productivity differed 
among nest substrates independent of height (see 
above), we used a nested ANOVA, with nest height as 
a main effect, and nest substrate (Artificial, Stump, and 
Tree) as the nested factor. There were no differences 
in productivity between either nest height (P = 0.50) 
or substrate (P = 0.08) in this analysis. 

There was no significant difference in nest success 
between Platform (58.6% successful) and Stump nests 
(68.5% successful; ica ='4.39, P= 0.11). Phe pite- 
portion of successful and unsuccessful nests varied 
among years Ga = 20.15, P = 0.03). The proportion 
of successful nests varied among years from 40.6% in 
1995 (all lakes, 32 nests) to 91.7% in 2001 (Sturgeon 
Lake only, 12 nests). 

Survival analysis was used to determine if longevity 
of nests varied among nest substrates. Because tests 
associated with survival analysis do not perform well 
at small sample sizes (StatSoft 2000), we excluded or 
pooled nest substrates with few observations. Thus, 
we compared Stump nests to Artificial nests (Utility 
pole and Platform). We were unable to get a good fit 
from several theoretical distributions. Annual survival 
of nests did not appear to vary between nest substrates 


TABLE |. Numbers of occupied unique nests and nest-years for Ospreys from 1991 to 2001 in the Kawartha Lakes. Nest-years 
was calculated by summing the numbers of years that nests of each substrate were occupied from 1991-2001. 


Number of 

Substrate individual nests 
Artificial 

Duck blind 5 

Utility pole 13 

Platform 19 

Total 37 
Natural 

Stump pa 

Tree 4 

Total 3] 


Nest-years (1991-2001) 


Number Percent of total 
| 4.2 
52 20.0 
85 32.7 
148 56.9 
96 36.9 
16 6.2 
1i2 43.1 


2005 


Proportion surviving 


—— Stump nests 
“-"" Artificial nests 


Age at beginning of interval (years) 


MARTIN, DE SOLLA, EWINS, AND BARKER: OSPREY NESTING 61 


—— Low nests 
=“"" High nests 


Age at beginning of interval (years) 


FIGURE 3. Proportion of nests surviving any given year in a) Stump and Artificial nests and b) High and Low nests. 


during the first three years of their use (Gehan’s Wil- 
coxon, P = 0.23), whereas after this age, Stump nests 
tended to have reduced survivorship compared to 
Artificial nests (Cox’s F-test, F 44, 44] = 1707? ]0:042: 
Figure 3a). Cox’s F-test (also known as log-rank) 
and Gehan’s Wilcoxon test emphasize the weight of 
later vs. earlier censored data, respectively (Fleming et 
al. 1987). Both Stump and Artificial nests surviving for 
6 years appeared to have relatively high survivorship 
after that point, although the sample size was small 
for these older nests. 

The comparison of longevity of High versus Low 
nests indicated that High nests had a greater longevity 
(Gehan’s Wilcoxon Test, P = 0.02). The survivorship 
of High nests was largely independent of age, where- 
as Low nests, which were primarily Stump nests, tend- 
ed to have reduced survivorship after three to five 
years (Figure 3b). Overall, High nests had greater 
survivorship than Low nests for all age classes. 


Discussion 

Ospreys breeding in the Kawartha Lakes region of 
south-central Ontario were found to nest successfully 
on low overwater flooded stumps. We found that, where 
stumps constituted the primary natural nest substrate, 
natural nests were more productive than nests built 
on artificial substrates. This contrasts with the findings 
of most other comparisons of natural and man-made 
Osprey nesting substrates. Ewins et al. (1995) found 
that natural nests, typically trees, were more likely to 
collapse and had lower productivity than artificial nest 


structures. Similarly, Westall (1983) found substan- 
tially higher productivity on artificial structures (1.47 
chicks per occupied nest) compared to natural nests 
(0.69) on Sanibel Island, Florida. Postupalsky (1977) 
also reported that nests on artificial substrates, typically 
man-made nesting platforms or utility poles, had high- 
er productivity than those on natural substrates. Witt 
(1996), however, found no differences in productivity 
from nests on man-made platforms versus trees in an 
Oregon population in which artificial nests comprised 
35% of total active nests. In many studies in the liter- 
ature however, natural nesting substrates constituted 
primarily trees, whereas in the current study most nat- 
ural nests were constructed on low flooded stumps. 
Stump nests in fact were almost twice as productive 
as tree nests in the current study, producing a mean of 
0.75 chicks per occupied nest more, annually. 
Despite the greater productivity of low stump nests, 
we found that their longevity was not as great as that 
of platform or utility pole nests. Survivorship of ele- 
vated nests appeared to be more independent of age 
compared to low nests, largely because of the perma- 
nency of the structure — utility poles and quadrupod 
platforms. Low nests consisted primarily of stumps, 
and occasionally abandoned duck blinds, frequently 
in disrepair. Thus, elevated nests tended to be avail- 
able every year, whereas the presence of stump nests 
was dependent upon sufficiently low water levels to 
prevent flooding or their destruction by ice action over 
the winter. We predict that the number of stumps 
available at Sturgeon and other flooded lakes within 


62 THE CANADIAN FIELD-NATURALIST 


the Kawartha Lake system will decline over the years 
as they degrade since further flooding of forested land 
is not expected. At that point Ospreys may switch to 
tree nesting or greater use of man-made platforms or 
utility poles. 

That Kawartha Lakes Ospreys nesting on flooded 
Overwater stumps were successful and productive is 
somewhat surprising, as Ospreys frequently nest on tall 
structures, presumably in part to avoid depredation. 
However, natural nests — mostly trees — are typically 
on shore, whereas the stump nests in our study were 
surrounded by water, far (usually > 100 m) from shore. 
Great horned owls (Budo virginianus) appeared to be 
an important predator of Ospreys in our study area; 
four nests were found with chicks depredated that were 
suspected to have been depredated by owls. Thus high 
nests may be equally or even more vulnerable than 
low nests to avian predators. Owl hunting patterns 
may emphasize forested and edge habitat, rather than 
over Open water where the stump nests are located. 
Another potentially important Osprey nest predator, 
raccoons (Procyon /oror), also present a greater risk 
to nearshore nests, and seem reluctant to swim far to 
access remote overwater nests (Poole 1989). Spring 
storms causing extreme wave action, waves from motor- 
boats, and sudden alterations in water levels through 
manipulating of dams, were occasionally suspected 
of flooding these low nests, resulting in their com- 
plete destruction, removal of eggs or chicks, or aban- 
donment (de Solla et al. 2003). Disturbance of nests 
by humans is another potential cause of nest abandon- 
ment or failure; nests located on overwater stumps 
are much more vulnerable to this impact than those 
high in trees. The Kawartha Lakes are popular recre- 
ational lakes, yet nests in open channels most acces- 
sible to humans appeared to suffer no greater failure 
than those in shallower areas with difficult boat access. 
The greater stability and possible lower risks of depre- 
dation conferred by overwater stump nests over tree 
nests appeared to account for their success and greater 
productivity in the current study. Certainly Ospreys 
in other locations have a propensity to nest in other 
low overwater sites, with channel navigational mark- 
ers being a notable example (Poole 1989). 

Many long-term programs have been implemented 
to monitor Osprey populations and improve recruit- 
ment, largely through the construction of artificial 
nesting sites (Barker 1988; Ewins 1996; Witt 1996). 
Most man-made nest structures consist of platforms 
erected upon a single pole on land, or upon a tripod 
or quadrupod in shallow water (Ewins 1994). The 
high proportion of stump nests at Sturgeon Lake was 
due to deliberate flooding caused by damming during 
the construction of the Trent-Severn waterway a centu- 
ry ago. Based upon the results of this study, artificial 
stump-like structures may also be suitable for Osprey 
nests, as long as they are built higher than the maximum 
water level during floods or annual fluctuations and 


Vol. 119 


are sufficiently far from shore. Artificial stump nests 
have the potential advantages of lower cost, lower 
maintenance, and fewer safety issues than taller struc- 
tures, and also allow for easier access by researchers 
to assess nest status and productivity of nests. 


Acknowledgments 

Funding for much of this work was supplied by 
The Ontario Ministry of Natural Resources and Envi- 
ronment Canada’s Great Lakes Action Plan. M. Bacro, 
L. Benner and G. Barrett provided assistance in the 
field. We are grateful for the Kawartha Lakes Friends 
of the Osprey Society for logistical support and for 
maintaining the artificial Osprey nest platforms. We 
also thank the Otonabee Region Conservation Author- 
ity, the former Ontario Hydro and the present Ontario 
Power Generation for their cooperation. 


Documents Cited (marked * in text) 

Barker, MLE. 1988. Osprey nesting survey, Lindsay District. 
Ontario Ministry of Natural Resources Report. 22 pages. 

Ewins, P. J. 1994. Artificial nest structures for Ospreys: a 
construction manual. Great Lakes Cleanup Fund, Environ- 
ment Canada, Canadian Wildlife Service, Toronto, Canada. 
41 pages. 


Literature Cited 

Castellanos, A., and A. Ortega-Rubio. 1995. Artificial nest- 
ing sites and Ospreys at Ojo de Liebre and Guerrero Negro 
Lagoons, Baja California Sur, Mexico. Journal of Field 
Ornithology 66: 117-127. 

de Solla, S. R., P. A. Martin, P. J. Ewins, and M. Barker. 
2003. Productivity and population trends of Ospreys in 
the Kawartha Lakes Region, Ontario, 1991-2002. Journal 
of Raptor Research 37: 306-313. 

Ewins, P. J. 1996. The use of artificial nest sites by an in- 
creasing population of Ospreys in the Canadian Great Lakes 
Basin. Pages 109-123 77 Raptors in human landscapes. 
Edited by D. Bird, D. E. Varland and J. J. Negro, Academic 
Press, London. 

Ewins, P. J., S. Postupalsky, T. Weise, and E. M. Addison. 
1995. Changes in the status, distribution and biology of 
Ospreys (Fandion haliaetus) breeding on Lake Huron. 
Pages 273-290 zz The Lake Huron Ecosystem: ecology, 
Fisheries and Management. Edited by M. Munawar, T. 
Edsall and J. Leach, SPD Academic Publishing, Amster- 
dam, The Netherlands. 

Fleming, T. R, D. P. Harrington, and M. O’Sullivan. 1987. 
Supremum versions of the log-rank and generalized Wil- 
coxon statistics. Journal of the American Statistical Asso- 
ciation 82: 312-320. 

Houston, C.S., and F. Scott. 1992. The effect of man-made 
platforms on osprey reproduction at Loon Lake, Saskat- 
chewan. Journal of Raptor Research 26: 152-158. 

Kaplan, E. L., and P. Meier. 1958. Nonparametric estima- 
tion from incomplete observations. Journal of the Ameri- 
can Statistical Association 53: 457-481. 

Martin, P. A., S. R. de Solla, and P. J. Ewins. 2003. Chlori- 
nated hydrocarbon contamination in Osprey eggs and nest- 
lings from the Canadian Great Lakes Basin, 1991-1995. 
Ecotoxicology 12: 209-224. 

Poole, A. F. 1989. Ospreys: a natural and unnatural history. 
Cambridge University Press, Cambridge, United Kingdom. 


2005 


Postupalsky, S. 1977. Artificial nesting platforms for Ospreys 
and bald eagles. Pages 35-45 7 Endangered birds: man- 
agement techniques for preserving endangered species. 
Edited by S.A. Temple, University Wisconsin Press, 
Madison, Wisconsin. 

StatSoft, Incorporated. 2000. STATISTICA for Windows 
[Computer program manual]. Tulsa, Oklahoma. 


MARTIN, DE SOLLA, EWINS, AND BARKER: OSPREY NESTING 63 


Westall, M. A. 1983. An Osprey population aided by nest 
structures on Sanibel Island, Florida. Pages 287-291 i 
Biology and management of bald eagles and Ospreys. Zdr- 
ed by D. M. Bird, Harpell Press, Ste. Anne de Bellevue, 
Quebec, Canada. 

Witt, J. W. 1996. Long-term population monitoring of Osprey 
along the Umpqua River in Western Oregon. Journal of 
Raptor Research 30: 62-69 


Received 19 November 2003 
Accepted 3 February 2005 


Effects of Wetland Creation on Breeding Season Bird Use in Boreal 
Eastern Ontario 


Davip A. Locky!“, J. CHRIS DavigEs2, and BARRY G. WARNER? 


'Biological Sciences Department, University of Alberta, Edmonton, Alberta T6G 2E9 Canada 

“Wetlands Research Centre and Department of Biology, University of Waterloo, Waterloo, Ontario N2L 3G1 Canada 
3Wildlife and Natural Heritage Section Ontario Ministry of Natural Resources, Peterborough, Ontario K9J 8M5 Canada 
4Present address: 11212 64" Street, Edmonton, Alberta TSW 4H3 Canada 


Locky, David A., J. Chris Davies, and Barry G. Warner. 2005. Effects of Wetland Creation on Breeding Season Bird Use in 
Boreal Eastern Ontario. Canadian Field-Naturalist 119(1): 64-75. 


Wetland construction has been an effective means of mitigating wetland habitat losses due to agricultural and other activities. 
However, the type, variety, and age of the habitats created are often critical components in the success of the wetland when 
the aim is to enhance the bird community. Hilliardton Marsh was constructed as a series of cells between 1993 and 1997 in 
boreal eastern Ontario to provide waterfowl habitat. We determined habitat change and monitored breeding-season bird use 
before construction and one year after the last cell was constructed. Wetland construction resulted in dramatic changes to 
the vegetation and bird communities. The area was transformed into a variety of wetland habitats, but primarily marsh, one 
of the rarest wetland types in boreal Ontario. Survey stations with moderate habitat change exhibited the greatest change in 
bird species richness. Total species richness increased 55% from 56 to 87 species, with obligate wetland birds increasing from 
3 to 26 species. Rare birds increased from 11 to 27 species, with most as obligate or facultative wetland birds, but also Peregrine 
Falcon (falco peregrinus). Bird abundance, as measured by the number of stations where a species was observed, increased 
significantly for obligate wetland birds. There were no significant losses of species from any bird group, as adjacent upland 
habitat was preserved. This short-term study has shown that construction of new wetland habitat in boreal eastern Ontario, 
especially marsh, can significantly increase the numbers of breeding-season birds, including rare species. However, long- 
term monitoring is required to ensure sustained success of wetland construction projects for birds. 


Key Words: boreal, breeding birds, constructed wetland, marsh, rare birds, upland birds, wetland birds, Ontario. 


Constructing wetlands to offset lost wetlands and fowl brood-rearing habitat, and secondarily as stag- 
maintain wildlife habitat has been widespread and ing habitat for waterfowl migrating to and from the 
relatively successful in North America (Whitman Hudson Bay and James Bay lowlands (Davies et al. 
1976; Leschisin et al. 1992; Creighton et al. 1997). 1996). It was constructed between 1993 and 1997 and 
In Ontario, most of the initiatives have occurred in comprises five cells contained within 8.6 km of dikes. 
the south where wetland losses have been extensive. ‘This site is managed by DU and water is drawn from 
Wetland creation has been less common in the bore- __ the adjacent Blanche River. 
al part of the province where wetland losses have not In May and June 1993, a pre-construction inven- 
been perceived to be important (Ritchie 1988). How- tory of the vegetation and bird communities at the 
ever, some boreal regions with glacio-lacustrine soils Hilliardton Marsh site was commissioned by the 
have seen significant numbers of wetlands lost to con- = Ontario Ministry of Natural Resources (OMNR) and 
version for agriculture. One such area in northeast- DU. Results from the 1993 inventory indicated that 
ern Ontario is the Little Clay Belt(LCB), where many __ the vegetation was predominantly upland and that only 
of the wetlands were drained for agriculture in the five species of wetland birds were observed on site 
early 1900s (Davies et al. 1996). (Davies et. al. 1996). Regionally rare birds were ob- — 

Marshes were more extensive before settlement in __ served on the site and included Upland Sandpiper (for 
the LCB and probably once supported a large popu- __ scientific names of birds see Table 3), Short-eared 
lation of waterfowl (Davies et al. 1996) (Figure 1). Owl, Eastern Meadowlark, and LeConte’s Sparrow. | 
Marsh habitat is particularly critical for wetland birds Preliminary post-construction vegetation (Gilbert — 
and is one of the rarest wetland types in the boreal _1997) and bird surveys (Huizer and Henshaw 1997*) 
region (National Wetlands Working Group 1988). To | were completed in 1997 before the wetland was fully 
address wetland losses in the LCB, Ducks Unlimited — constructed. Results indicated that shallow water wet- 
Canada (DU) and its partners have initiated a num- _land and marsh habitat was created and that waterfowl 
ber of wetland creation projects, including Hilliardton and other wetland birds were attracted to the wetland — 
Marsh in the Hilliardton Marsh Provincial Wildlife | (Huizer and Henshaw 1997*). 


Area (HMPWA) (Figure 1). This constructed wetland Few studies have examined the breeding bird com- 
is 209ha and one of the largest projects of its kind in —_munities in the Clay Belt of northern Ontario (Smith 
Ontario. The wetland was created primarily for water- 1957; Erksine 1977), which is an important area from 


64 


2005 Locky, DAVIES, AND WARNER: 


nn 


290. 


(foreground) 


— 
et pe 


EFFECTS OF WETLAND CREATION 65 


Por 


at 


Control Stations | 


FicureE 1. Hilliardton Marsh with 24 wetland bird survey stations on the dikes. Inset image shows five of six control 


stations outside and northeast of the wetland. 


the perspective of marking the northern, eastern, and 
southern limits of the ranges of several bird species 
(Smith 1957). The purpose of this study was to deter- 
mine the effects on the breeding-season bird commu- 
nity of complete construction of Hilliardton Marsh 
one year after completion and relate any changes to 
habitat transformation. The cells were aged from one 
to five years at the time of the survey. Our intent was 
not to provide a definitive assessment of wetland con- 
struction on breeding-season birds, but to illustrate the 
initial impacts of wetland construction on the bird 
community. Our specific objectives were to: (1) quan- 
tify habitat change in area flooded for the wetland 
and at the survey stations, (2) compare total and bird 
group species richness before and after wetland con- 
struction, (3) compare total and bird group species 
abundance before and after wetland construction, (4) 
determine changes in total and bird group rare spe- 
cies between years, and (5) examine the relationship 
| between degree of habitat change and bird species 
| richness between years. 

We hypothesized that post-construction bird surveys 
would be dominated by wetland dependent species 
} at stations with high habitat change, whereas survey 
} Stations with little or no habitat change would con- 
| tinue to be dominated by upland species. Stations with 
| moderate habitat change would have the highest num- 


ber of species and abundances. We hypothesized that 
rare species would increase with the addition of wet- 
land habitat and we expected no change in any vari- 
ables at the control stations. 


Study Area 

Hilliardton Marsh Provincial Wildlife Area is 20 km 
north of New Liskeard, Ontario, Canada (47°46'N; 
79°42'W; 190 m a.s.1.). The topography is relatively 
low relief and the glacio-lacustrine soils are clay-rich 
(Dredge and Cowan 1989), which together have con- 
tributed to poor drainage. The site lies in the greater 
Mid-Boreal Wetland Region where peatlands are the 
dominant wetland types (National Wetlands Working 
Group 1988) and approximately 4% is marsh (Riley 
1988). 

The LCB has cropland and pasture, second-growth 
mixed hardwood forest, and various types of natural 
wetlands that include mixed deciduous and coniferous 
swamp, fen, bog, and tall shrub swamp (Davies et al. 
1996). Prior to construction, Hilliardton Marsh was 
agricultural land in various states of use and abandon- 
ment, with some second-growth Trembling Aspen 
(Populus tremuloides) forest. Approximately 20% of 
the area was shrub swamp dominated by Speckled 
Alder (A/nus incana ssp. rugosa) and willows (Sa/ix 
spp.) (Davies et al.1996). 


66 THE CANADIAN FIELD-NATURALIST 


Methods 
Habitat Survey 

Pre-construction habitat was delineated using a 
site plan (Ducks Unlimited 1986*) and post-construc- 
tion habitat with 1998 aerial photographs (1:5000) 
and field reconnaissance. Description of the wetland 
classes followed the Canadian Wetland Classification 
System (National Wetlands Working Group 1997) 
and vegetation communities were defined by dominant 
vegetation forms following the Northern Ontario 
Wetland Evaluation (Ontario Ministry of Natural 
Resources 1994). The aereal coverage for each of the 
vegetation communities was calculated with a light 
planimeter for 1993 and 1998, and the extent of habitat 
change after flooding was determined. Habitat change 
within each of the 30 200 m radius (12.6 ha) point- 
count stations was also calculated. All of the wetland 
survey stations except two contained primarily upland 
habitat before flooding. Habitat change was therefore 
considered a general indicator of new wetland habitat. 
Stations were classified into three categories based on 
percent habitat change as follows: little or no change 
(O-19%, O-2.4 ha), moderate change (20-54%, 2.5- 
6.8 ha), and high change (55-100%, 6.9-12.56 ha). 


Bird Surveys 

Thirty 200 m radius avian point-count stations were 
established as repeatable survey points in 1993 
(Huizer and Henshaw 1997) (Figure 1). Twenty-four 
of the wetland survey stations were placed 400 m 
apart (centre to centre) on the proposed wetland im- 
poundment berms and were used to detect changes 
in the bird community related to wetland construc- 
tion. Six control stations were located 400 m outside 
the dikes away from the flooding and were used to 
detect changes in species composition not associated 
with the flooding event (e.g., weather) [Inset image 
in Figure 1]. 

To provide a bird community data set comparable 
to that from 1993, we completed bird surveys at the 
same stations, during the same survey period, and 
for the same duration as the 1993 surveys. Different 
observers were used in 1993 and 1998, but the survey 
standardization minimized the probability of bird 
diversity changes between years being attributed to 
unequal sample effort (Elphick 1997). Stations were 
surveyed between sunrise and 10:00 am when there 
was good visibility, no precipitation, and little wind. 
Upon arrival at the station the surveyor faced north, 
waited five minutes, and then recorded all bird move- 
ments and auditory calls during a 10-minute interval 
(Canadian Wildlife Service 1997*). Pre-construction 
point-counts were conducted 22-24 May and 11-13 
June 1993. Post-construction surveys were conduct- 
ed 23-25 May and 18-20 June 1998. Birds were also 
noted outside of the survey circles in the study area 
during the survey period in both years to determine 
total species richness and composition. 


Vol. 119 


Bird Groups and Species Classification 

Bird species abundance (e.g., common, rare) and 
status (e.g., breeder, migrant) within the LCB region 
were determined according to a regional bird list 
(Temiskaming Field Naturalists 1994*) and the North- 
ern Ontario Wetland Evaluation System (Ontario 
Ministry of Natural Resources 1994). Species were 
considered breeding on the site if there was a singing 
male on territory; a pair was observed, individuals 
were carrying food, or there was agitated behaviour 
(territoriality) (Huizer and Henshaw 1997*). Species 
recorded for counts during the May period were only 
considered breeding on the site if they were within 
their traditional breeding range, whereas species out- 
side their known range were considered migrants. 
Species classified as migrants were removed from the 
breeding-bird data-sets in order to compare breed- 
ing-season birds within the LCB region. However, 
all species in the area during surveys were recorded 
by including birds observed outside of survey plots. 
The May and June bird surveys were combined to in- 
clude both early and late breeding species (approxi- 
mately 15 May — 15 July). 

We classified bird species into five groups based on 
wetland/aquatic habitat dependency as determined 
and further modified by life history descriptions in 
Cadman et al. (1987) to better reflect boreal bird com- 
munities as follows: 

I. Obligate Wetland (found greater than 99% in wet- 
lands); 
II. Facultative Wetland (57-99%, generally found in 
or near wetlands); 
II. Facultative (34-56%, occurs frequently in wet- 
lands, but wetlands are not essential); 
IV. Facultative Upland (1-33%, occasional or no use 
of wetlands); and 
V. Upland (found greater than 99% in uplands). 
The assignment of species groups or rankings to — 
species and the results of associated statistical analy- 
ses have been a matter of discussion among experts 
(Simberloff and Dayan 1991). In particular, assign- 
ment of scarce species to categories, however defined, 
may be error-prone. However, our intent in using bird 
groups was to consider and monitor the bird commu- 
nity as a whole, with minor differences in individual 
rankings having little impact on the final analysis 
(c.f. Croonquist and Brooks 1991). Bird nomenclature 
in this study follows the American Ornithologist Union 
(AOU) 7" Edition Checklist (AOU 1998), and sup- 
plements 42 (AOU 2000), 43 (AOU 2002), 44 (AOU 
2003), and 45 (AOU 2004). 


Data Analysis 

We assumed that the point-count circles were large 
enough (200 m radius, 12.6 ha in area) to include all 
or portions of territories for many breeding-season 
bird species and that the total coverage was repre- 
sentative of all habitat types measured at Hilliardton 


2005 


Marsh. The stations were 400 m apart, much farther 
than the minimum 250 m suggested by Freemark and 
Rogers (1995) and Ralph et al. (1995) to limit double 
counting. However, although station edges were touch- 
ing and some independence in bird observations may 
be compromised, the temporal scale was relatively 
small and the change in species richness and presence 
at stations was of most interest to us in this study (see 
Bibby et al. 1993). 

The calculation of bird abundance directly from 
point-count data is problematic (Bibby et al. 1993; 
Ralph et al. 1995). This is especially true with large 
survey circles as used in this survey, as the detect- 
ability of birds varies greatly among different habitats 
(Hutto et al. 1986), among bird species (Ralph et al. 
1995), and distance from observers. Instead, we used 
the number of stations at which a species was ob- 
served each year as a proxy for species abundance 
(station/species/year). We assumed that if the actual 
abundance of a species increased, the probability that 
at least one bird is observed at a station would increase. 
Although there is potential to artificially inflate abun- 
dance due to the possibility of several stations equat- 
ing the presence of only a single individual of a spe- 
cies, there is also a higher probability that a species will 
be present at a station and thus the expected number 
of stations at which that species is observed will also 
increase (Debinsky and Humphrey 1997). 

Species counts were categorized by bird group and 
year, and by degree of habitat change and year. For 
statistical analysis, we assumed that bird observation 
stations and observations were independent of each 
other and utilized time-series statistical analyses 
(Hurlbert 1984). Wilcoxon’s Rank Sign tests were 
used, as the untransformed and transformed data 
failed Levene’s Test of Equality (SPSS Inc. 2002). 
This test was used to determine significant changes in 
mean numbers of total species and species/station for 
the wetland stations and control stations, changes 
between years for number of stations/species, a proxy 
for abundance, and changes between years of species/ 
station by low, moderate, and high habitat change. 
The McNemar Test for Significance of Changes (Con- 
over 1980) was used to test for the significance of 
between-year (1993, 1998) changes in the number of 
Stations at which a particular species was observed. 
All data analyses were completed using SPSS ver. 
11.5.1 (SPSS Inc. 2002), except for the McNemar Test 
which was programmed using Microsoft Excel®. 

This study was somewhat constrained by the survey 
design used for the original 1993 study, and presented 
a number of limitations on the interpretation of our 
results. The Canadian Wildlife Service point count 
survey protocol was not specifically designed for open 
wetlands, for which the employment of broadcast bird 
calls to illicit responses from more secretive wetland 
birds is often used; wetland birds are often less evenly 
distributed than upland species, increasing the diffi- 


Locky, DAVIES, AND WARNER: EFFECTS OF WETLAND CREATION 67 


culty of sampling efforts (Bell et al. 1973). Because 
only two survey stations were not peripheral (i.e., did 
not include edge), diversity may be unrepresentatively 
high for wetland birds, and under sampled in other 
stations (see Erskine 1977). The use of six control sta- 
tions may not be enough to accurately detect changes, 
as even if habitats were uniform, changes detected may 
not be truly representative for the area. With respect 
to the May count dates in low boreal habitats, some 
insectivorous birds may be under-represented due to 
not having returned in breeding numbers. Addition- 
ally, some of the rare species may be vagrants outside 
of their normal breeding range. Despite these limita- 
tions, we feel that the data contributes much needed 
information on not only breeding season birds in the 
Clay Belt where there have been few studies, but on 
the effects to birds of creating rare wetland habitat in 
a region with high wetland losses. 


Results 
Habitat Change 

Habitat change was dramatic after the construc- 
tion of Hilliardton Marsh. The agricultural fields that 
encompassed 71% of the site (145 ha) were largely 
submerged and all of the Trembling Aspen forest 
was flooded (13% or 27 ha) (Table 1). The new wet- 
land consisted of 54% marsh (112 ha), wetland with 
shallow water and dominated by emergent aquatic 
graminiods, 29% deciduous swamp (61 ha), wooded 
wetland dominated by either trees or shrubs, and 13% 
shallow water wetland (27 ha), wetland with standing 
or flowing water less than 2 m in mid-summer, and 
dominated by floating aquatic macrophytes (National 
Wetlands Working Group 1997). Aerial photographs 
taken in 1998 indicated that a large proportion of the 
marsh exhibited hemi-marsh habitat (50:50 open water/ 
vegetation), which is attractive to aquatic birds (Wel- 
ler 1994). 

Habitat change at the survey stations was also sig- 
nificant, with four stations in the high change category 
(55-100%), 20 stations in the moderate change catego- 
ry (20-54%), and the six control stations in little/no 
change category (0-19%). Fifteen of the 24 stations 
that were flooded as a result of wetland construction 
exhibited habitat change of at least 50%. 


Total Bird Species Richness 

After wetland construction the total bird species 
richness on site increased 34% from 72 species in 1993 
to 109 species in 1998. The 109 species represent 49% 
of the 222 birds that are known to occur on the LCB 
at any time as year-round residents, migrants, breeders, 
or winter residents (Temiskaming Field Naturalists 
1994*) (Tables 2 and 3). At the HMPWA, 56 breed- 
ing species were observed, which reflects 36% of the 
156 breeding species found in the in the LCB. This 
number increased to 87 species in 1998, or 56% of 
total observed breeding species in the LCB. 


68 THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |. Summary of habitat type and area before and after construction of Hilliardton Marsh. Numbers in brackets are 


percent of total wetland. 


1993 
Habitat Type Area 
Early Successional Poplar/Alder Forest 14.0 
Deciduous Woods 9.0 
Early Successional White Birch/Poplar Fores 4.0 
Active Agricultural Field 3:0 
Fallow/Hay Agricultural Field 1395 
Alder/Willow/Poplar Swamp 37D 
Shallow Water Wetland 0.0 
Marsh 0.0 
Dead Tree/Shrub Swamp 0.0 
Total 209.0 


Survey Stations and Bird Groups 

Before construction of Hilliardton Marsh in 1993, 
most birds observed at survey stations were upland 
(30%), facultative upland (25%) and facultative spe- 
cies (25%) (Tables 2 and 3). Only 14% were faculta- 
tive wetland species and 5% were obligate wetland 
species. One year after wetland construction (1998), 
30% of the survey station birds were obligate wetland 
species, with 22% each as facultative and upland 
species, 17% as facultative upland, and 9% as facul- 
tative wetland. 

There were significant differences between years 
for overall richness at the wetland survey stations 
(P < 0.001), and the average number of species/sta- 
tion increased from 12.6 in 1993 to 20.7 in 1998. These 
differences, however, were not constant across the 
groups (Figure 2). Birds in three groups increased 
significantly in richness including obligate wetland 
species, 0.5 to 7.0 species/station (P < 0.001), facul- 
tative wetland species, 1.5 to 2.0 species/station (P = 
0.011), and facultative upland species, 3.5 to 4.4 
species/station (P = 0.026). There were no significant 
differences between years for overall richness or 
number of species/station for total or grouped birds 
at the control stations. 


1998 % Change 

(ha) Area (ha) 1993-1998 
(7) 0.0 (0) (-7) 
(4) 0.0 (0) (-4) 
(2) 0.0 (0) (-2) 
(2) 0.0 (0) (-2) 
(67) 9.0 (4) (-63) 
(18) 34.0 (16) (-2) 
(0) 27.0 (43) (+13) 
(0) 112.0 (55) (+54) 
(0) 27.0 (13) (+13) 
(100) 209.0 (100) (0) 


Small increases in numbers of stations/species, a 
proxy for abundance, were observed in all five groups 
(Table 3), but significant differences were only evi- 
dent in the obligate wetland group (P < 0.001). This 
group increased from 0.5 to 8.0 stations/species. 
Regionally rare Pied-billed Grebe, American Wigeon, 
and Northern Shoveler had particularly high relative 
increases in observations between years (Table 3). 
The provincially rare Yellow Rail was counted at one 
station during the 1998 survey, but individuals were 
heard calling in at least two other locations within 
the marsh during Twenty-one species had significant 
differences (P = 0.039 to P < 0.001) in station counts 
between 1993 and 1998, most of these as increases. 
Twelve species (57%) were obligate wetland, two were 
facultative wetland, four were facultative, and three 
were facultative upland species. All obligate wetland 
species had positive changes between years. Those 
species with losses included Wilson’s Warbler (facul- 
tative wetland) (P = 0.002), Alder Flycatcher (faculta- 
tive) (P = 0.012), Ruby-crowned Kinglet (facultative) 
(P = 0.008), and Mourning Warbler (facultative) (P = 
0.039). Red-winged Blackbirds (facultative wetland) 
were recorded in 23 wetland stations and one control 
station in 1998; there were only two station observa- 


TABLE 2. Summary of bird species richness by bird group at site and survey stations with percentages (in brackets) before 
and after wetland construction. Comparisons with total possible species in region and possible breeders in region at the bot- 
tom. Obligate Wetland (found greater than 99% in wetlands), Facultative Wetland (57-99%, generally found in or near wet- 
lands), Facultative (34-56%, occurs frequently in wetlands, but wetlands are not essential), Facultative Upland (1-33%, 
occasional or no use of wetlands), and Upland (found greater than 99% in uplands). 


Bird Possible Possible 

Group in Region Breeding 

Obligate Wetland 72 (32) 40 (26) 5) 
Facultative Wetland 16 (7) 1] (7) 8 
Facultative 30 (14) 26 (17) 16 
Facultative Upland 33 (15) 26 (17) 16 
Upland 71 (32) 33 (34) 27 
Total bee. '; (O00) 156 (100) IZ 


% of Total Possible Species in the Region 
% of Possible Breeding Species in the Region 


Site 


Before (1993) After (1998) 
Survey Site Survey 

(7) 3 (5) 34 (31) 26° ~*(80) 
(11) 8 (14) 10 (9) 8 (9) 
(22) 14 (25) 20 (18) 19" (22) 
(22) 14 (25) 20 (18) 15 (17) 
(38) Le (30) 25 (23) LG ivy shed) 
(100) 56 (100) 109 = (100) 87 (100) 
(32) (25) (49) (39) 
(46) (36) (70) (56) 


2005 


Locky, DAVIES, AND WARNER: EFFECTS OF WETLAND CREATION 


69 


TABLE 3. Bird species observed at the site and within survey stations before and after construction of Hilliardton Marsh. 
Stations column lists the number of stations in which a species was observed, which is a proxy for abundance. Asterisk 


denotes species observed within and outside of survey stations. See text for explanation of bird groups. 


tions in 1993 (Table 3). Presence of the regionally 
rare LeConte’s Sparrow remained constant at four 
stations between 1993 and 1998. 


Habitat Change and Bird Groups 
Overall numbers of species/station increased 


Bird Species eT Bird Before After 
Common Name Scientific Name Status Group Site Stations Site Stations 
Canada Goose Branta canadensis CM ‘i 3 
Wood Duck Aix sponsa CB . 6 
American Wigeon Anas americana CB + 1] 
American Black Duck Anas rubripes CB * l 
Mallard Duck Anas platyrhynchos CB . 2 16 
Blue-winged Teal Anas discors (i i 14 
Northern Shoveler Anas clypeata UB - 10 
Northern Pintail Anas acuta CM * 
Green-winged Teal Anas crecca CB # J 
Ring-necked Duck Aythya collaris CB 53 3 
Common Goldeneye Bucephata clangula CB - 8 
Hooded Merganser Lophodytes cucullatus CB * l 
Common Merganser Mergus merganser CB " l 
Pied-billed Grebe Podilymbus podiceps CB “3 20 
Horned Grebe Podiceps auritus RM Z * 2 
Double-crested Commorant Phalacrocorax auritus RM = $ 
American Bittern Botaurus lentiginosus CB > é 12 
Great Blue Heron Ardea herodtas CB » 2 | 
Yellow Rail Coturnicops noveboracensis RB Sp a l 
Virginia Rail Rallus limicola RB ra - 3 
Sora Porzana carolina CB © $ 19 
American Coot Fulica americana CB . G2 
Sandhill Crane Grus canadensis RB ? 
Lesser Yellowlegs Tringa flavipes CM ‘i 
Spotted Sandpiper Acitis maculartus CB zi 3 
Wilson’s Snipe Gallinago delicata CB - 2 5 26 
American Woodcock Scolopax minor CB iy 
Ring-billed Gull Larus delawarensis CB a, | 
Black Tern Chlidonias niger RB = 3 
Belted Kingfisher Ceryle alcyon CB ai 
Marsh Wren Cistothorus palustris RB ‘3 
Northern Waterthrush Seirus noveboracensis UB a 3 . 8 
Swamp Sparrow Melospiza georgiana CB - 9 * 20 
Total in Group I; Brackets — total stations per species for group a 3 (14) 34 =. 26 (210) 
Bird Species Mbnal Bird Before After 

Common Name Scientific Name Status Group Site Stations Site Stations 
Upland Sandpiper Bartramia longicauda RB ss 

_ Northern Harrier Circus cyaneus CB * | ‘ | 
Great Gray Owl Strix nebulosa RB a = I 
Yellow-bellied Flycatcher Empidonax flaviventris CB S . 4 - aha 
Winter Wren Troglodytes troglodytes UB 3 ‘3 gi z 1 
Northern Parula Warbler Parula americana R S + 
Palm Warbler Dendroica palmarum CM is ~ 
Common Yellowthroat Geothlypis trichas CB iS zt 19 - 22 
Wilson’s Warbler Wilsonia pusilla CM 3 = 11 . | 
Le Conte’s Sparrow Ammodramus leconteit RB 2 * 4 f 4 
Red-winged Blackbird Agelaius phoeniceus CB x 2 : 24 
Brewer’s Blackbird Euphagus cyanocephalus CB - i 
Total in Group II; Brackets — total stations per species for group 9 8 (44) 10 8 (58) 


between years in similar magnitude at stations with 


70 THE CANADIAN FIELD-NATURALIST Vol. 119 
TABLE 3. continued. 

Bird Species aie Bint Before After 
Common Name Scientific Name Status Group Site Stations Site Stations 
Short-eared Owl Asio flammeus CB - 
Ruby-throated hummingbird = Avchilochus colubris CB 2 : 
Alder Flycatcher Emptidonax alnorum CB ss 19 s 10 
Gray Jay Perisoreus canadensis Gis . 5 * l 
Common Raven Corvus Corax CR # * 4 
Tree Swallow Tachycineta bicolor CB * 12 
Cliff Swallow Petrochilidon pyrrhonota CB * 1 
Boreal Chickadee Poecile hudsonica RW * 2 
Ruby-crowned Kinglet Regulus calendula CB “3 12 ss 
Veery Catharus fuscescens CB $ s iS ‘3 i 
Swainson’s Thrush Catharus ustulatus CB Ss 5 3 3 
Hermit Thrush Catharus guttatus CB 2 - 4 _ 4 
Gray Catbird Dumetella carolinensis UB = i 2 
Nashville Warbler Vermivora ruficapilla CB i 18 is 15 
Yellow Warbler Dendroica petechia CB 7 4 sg 7 
Black-throated Blue Warbler = Dendroica caerulescens RB 3 1 
Yellow-rumped Warbler Dendroica coronata CB - 6 - 2 
Connecticut Warbler Oporornis agilis RB is 
Mourning Warbler Oporornis philadelphia CB ss 15 - iz 
Song Sparrow Melospiza melodia CB - 14 % 18 
Lincoln’s Sparrow Melospiza lincolnit CB ‘ a ‘g 3 
Bobolink Dolichonyx oryzivorus CB _ 10 2 6 
Total in Group III; Brackets — total stations per species for group 17 14 (131) 20 ASCE 

Bird Species Abang Bird Before After 
Common Name Scientific Name Status Group Site Stations Site Stations 
Peregrine Falcon Falco peregrinus RB a 
Killdeer Charadrius vociferus CB 4 a 1 
Northern Flicker Colaptes auratus CB z 8 z 5 
Pileated Woodpecker Dryocopus pileatus CR & - 4 
Least Flycatcher Empidonax minimus CB - 14 : 11 
Eastern Kingbird Empidonax tyrannus CB Z 10 
Warbling Vireo Vireo gilvus R : a 
American Crow Corvus brachyrhynchos CR 2 = 10 
Barn Swallow Hirundo rustica CB = . 1 S p 
Wood Thrush Hylocichla mustelina RB "A i’ 1 
American Robin Turdus migratorius CB = = 19 22 
Magnolia Warbler Dendroica magnolia CB 42 : 4 ss 1 
Black-throated Green Warbler Dendroica virens RB = e 
Black-and-white Warbler Mniotlta varia CB 3 « 1] . 6 
Cedar Waxwing Bombycilla cedrorum CB oo 6 l i 
Savannah Sparrow Passerculus sandwichensis CB % 14 5 14 
White-throated Sparrow Zonotrichia albicollis CB 2 23 * 18 
Common Grackle Quiscalus quiscula CB e 3 3 22 
Baltimore Oriole Icterus galbula UB 2 2 
Purple Finch Carpodacus purpureus CB a | is 
White-winged Crossbill Loxta leucoptera UW ss 
Total in Group IV; Brackets — total stations per species for group 16 14 (103) 20) AS ise) 


moderate habitat change (12.5 to 21.9) and high 
habitat change (12.3 to 23.5), but the results were 
only significant for the former (P < 0.001). Stations 
with no habitat change had 13.3 and 14.0 species/ 
station for each year. The high variability in the high 
habitat change data suggests the sample size of four 
was too small for significant results (P = 0.068). By 
bird group, the distribution of species was somewhat 


more even and less variable in stations with moderate 
habitat change than in the other groups. However, 
the results were not significant (Figure 3). 


Rare Spectes 

Twenty-seven rare species were observed on site 
during the 1998 survey, more than double the 11 rare 
species observed in 1993 (Table 4). The most notable 
new rare species were the nationally threatened Pere- 


2005 


TABLE 3. continued. 


Locky, DAVIES, AND WARNER: EFFECTS OF WETLAND CREATION 71 


Bird Species 


Common Name Scientific Name 


Ruffed Grouse 
Broad-winged Hawk 
American Kestrel 
Merlin 

Mourning Dove 
Black-billed Cuckoo 
Great Horned Owl 
Yellow-bellied Sapsucker 
Downy Woodpecker 
Hairy Woodpecker 
Eastern Wood Pewee 


Bonasa umbellus 
Buteo platypterus 
Falco sparvertus 
Falco columbarius 
Zenatda Macroura 
Coccyzus erythropthalmus 
Bubo virginianus 
SPAYrapicus Vartus 
Picoides pubescens 
Picoides villosus 
Contopus virens 


Blue-headed Vireo Vireo solitarius 
Philadelphia Vireo Vireo philadelphicus 
Red-eyed Vireo Vireo olivaceus 

Blue Jay Cyanocitta cristata 
Horned Lark Evremophila alpestris 


Black-capped Chickadee 
Red-breasted Nuthatch 
Brown Thrasher 
European Starling 
Tennessee Warbler 
Chestnut-sided Warbler 


Poecile atricapillus 
Sitta canadensis 
Toxostoma rufum 
Sturnus vulgaris 
Vermivora peregrina 
Dendroica pensylvanica 


Cape May Warbler Dendroica tigrina 
American Redstart Setophaga ruticilla 
Ovenbird Seirus aurocapillus 
Chipping Sparrow Spizella passerina 


White-crowned Sparrow 
Northern Cardinal 
Rose-breasted Grosbeak 
Eastern Meadowlark 
Pine Siskin 

American Goldfinch 
Evening Grosbeak 
House Sparrow 


Zonotrichia leucophrys 
Cardinalis cardinalis 
Pheucticus ludovicianus 
Sturnella magna 

Carduelis pinus 

Carduelis tristis 
Coccothraustes vespertinus 
Passer domesticus 


Total Group V; Brackets - total stations per species for group 


Abun/ 
Status 


@) 

w 

* % 
¥ ¥ * *® * * 


Q 
oO 
= 
oO’ 


(eteke! 
w v2) 
Upland 
* 
eS 


(@ 
w 
* 


Q 
ee) 
* * & % 


Before After 


Bird 


Group Stations 


l 


Site Stations Site 


3 
l 


—_ 
xe KE  E KF HF HE KH F 


i 
* + & 
~) 


i ee ates 
ae 
—_—_— — 


5 s 9 


CR : l 


p 4 | 17 (85) 2S 19 (97) 


Abundance and status codes: CB — common breeder, RM — rare migrant, CM — common migrant, UB — uncommon breeder, 
C — common, R — rare, RB — rare breeder, RW — rare in winter, VB — very rare breeder, VM — very rare migrant, UW — 


uncommon in winter, V — very rare, CR — common resident. 


grine Falcon and four provincially significant spe- 
cies, including Horned Grebe, American Coot, Yellow 
Rail, and Black Tern. Most of the rare species ob- 
served in 1998 had some degree of wetland depend- 
ency, with 14 as obligate wetland and four as facul- 
tative wetland. In contrast, only two of the 11 rare 
species observed in 1993 were provincially signifi- 
cant: Short-eared and Great Gray Owls. Five rare 
species surveyed in 1993 were not observed during the 
post-constructions surveys and included Short-eared 
Owl, Upland Sandpiper, Connecticut Warbler, Wood 
Thrush, and Great Gray Ow] (Table 4). 


Discussion 
Habitat Change and the Bird Community 

Habitat heterogeneity can be an important element 
in attracting a wide variety of bird species, especially 
| in wetlands (Creighton et al. 1997). Wetland complex- 
es of varying water depths and flooding duration often 


lead to the most comprehensive array of habitat types 
for birds (Murkin et al. 1997). Habitat change at 
Hilliardton Marsh was not only dramatic but resulted 
in an overall increase in habitat types. In 1998, 54% 
of Hilliardton Marsh was marsh habitat one to five 
years old. Increased vegetation diversity, percent cov- 
er, and structure attract a greater diversity of birds 
compared to less vegetated marshes (Whitman 1976; 
Kantrud and Stewart 1984; Delphey and Dinsmore 
1993; Weller 1994; VanRees-Siewert and Dinsmore 
1996). Many edge habitats were created and these are 
prime habitat for many bird species (Cyr et al.1995). 
Where water meets land, i.e., riparian areas, insectiv- 
orous bird abundance is often the greatest (Iwata et al. 
2003). Survey stations with moderate habitat change 
tended to have the greatest diversity of habitats, many 
with hemi-marsh vegetation development, and in turn 
had significant increases in total species richness. 


2 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE. 4. Rare birds by status and bird group at Hilliardton Marsh Provincial Wildlife Area before (1993) and after wetland 
construction (1998). Rarity determined by Temiskaming Field Naturalists (1994*) for upland and wetland birds, Ontario 
Ministry of Natural Resources (1994) for wetland birds, and Environment Canada (2004*) for threatened birds. See text for 
explanation of bird groups. 


Status Bird Group Species Before After 
Threatened Facultative Upland Peregrine Falcon ** 
Obligate Wetland Horned Grebe * 
Obligate Wetland Yellow Rail * 
Provincially Obligate Wetland American Coot - 
Significant Obligate Wetland Black Tern a 
Facultative Wetland Great Gray Owl si 
Facultative Short-eared Owl * 
Obligate Wetland American Wigeon * 
Obligate Wetland Northern Shoveler sg 
Obligate Wetland Pied-billed Grebe * 
Obligate Wetland Double-crested Cormorant = 
Obligate Wetland Virginia Rail * 
Obligate Wetland Sora 3 
Regionally Obligate Wetland Sandhill Crane = 
Significant Obligate Wetland Marsh Wren * 
Facultative Wetland Upland Sandpiper - 
Facultative Wetland Le Conte’s Sparrow 2 * 
Facultative Connecticut Warbler e 
Facultative Upland Warbling Vireo - * 
Facultative Upland Wood Thrush . 
Upland Horned Lark . * 
Upland Eastern Meadowlark ? a 
Obligate Wetland Canada Goose ‘i 
Obligate Wetland Northern Waterthrush s ‘a 
Locally Facultative Wetland Winter Wren * = 
Significant Facultative Gray Catbird * 
Facultative Upland Baltimore Oriole * 
* 


Upland 


Brown Thrasher 


Constructed wetlands may evolve towards the 
ecological characteristics of natural wetlands within 
a few years (Sistani et al. 1999), and wetland age has 
been positively correlated with increases in native wet- 
land plant species diversity and richness (Reinartz 
and Warne 1993). Compared to younger wetlands with 
less vegetation development, wetlands four years of 
age have been found to have higher wetland avian spe- 
cies richness (VanRees-Siewert and Dinsmore 1996; 
Wan et al. 2001). This corresponds to the average 
wetland cell age at Hilliardton Marsh of 3.5 years 
with a range of 1-5 years in 1998. Smaller areas with 
homogeneous stands of Cattail (Zipha latifolia) (with 
some open water) are critical for attracting species 
such as Marsh Wren (Verner and Engelson 1970), and 
this rare regional breeder was first seen at Hilliardton 
Marsh in 1998. However, vegetation communities are 
dynamic and this study captured only the initial ef- 
fects of wetland vegetation development on the bird 
community. In new impoundments, plant succession 
tends to advance to stable, rooted aquatic plants and 
rapidly increasing invertebrate populations have been 
found to stabilize within two years. Thus, the initial 
conditions that proved favourable for many species of 


waterbirds diminishes (Whitman 1976). Unless the 
vegetation communities are maintained for a diversity 
of habitat types by managers, the bird community at 
Hilliardton Marsh will continue to change, especially 
for obligate wetland species. For example, increased 
growth of Cattail between 1998 and 1999 was docu- 
mented and would attract certain birds; e.g., Marsh 
Wren and Red-Winged Blackbird, but displace others 
that favour more open habitats, e.g., diving ducks. 
Wooded wetland habitats such as treed swamps are 
used by some waterfowl for nesting or cover (Cad- 
man et al. 1987; Merendino et al. 1995), including 
Common Goldeneye, Ring-necked Duck, Hooded 
Merganser, and Wood Duck (Table 3). The treed 
swamps at Hilliardton Marsh are temporary habitat, 
as the dominant tree 1s Trembling Aspen, a non-wet- 
land species. Although these trees were alive one year 
after the Hilliardton Marsh was constructed, the ex- 
tended flooding would eventually kill the trees leaving 
a dead-tree swamp (c.f. Ontario Ministry of Natural 
Resources 1994). That habitat structure will even- 
tually collapse with high water levels and inevitably 
become marsh or shallow water habitat that is less 
attractive to birds with wooded-wetland affinities. 


2005 


Mam Before Construction 
(205 After Construction 


Species Richness 


ObIWet FacWet_ Fac Fac Up Upland 


Bird Groups 


FIGURE 2. Species richness by bird group before and after 
wetland construction with 95% confidence interval 
error bars. OB] Wet = Obligate Wetland, Fac Wet = 
Facultative Wetland, Fac = Facultative, Fac Up = 
Facultative Upland. See text for explanation of bird 
groups. 


Rare Habitat, Rare Species 

Marsh habitat is the rarest wetland habitat in north- 
ern Ontario (National Wetlands Working Group 1988; 
Riley 1988). Bird species from the southern boreal 
region of Ontario and Quebec that inhabit open water 
marsh (and swamp) habitats are limited by the occur- 
rence of these habitats (Cadman et al. 1987; Gauthier 
and Aubry 1996). Similar studies on the effects of 
restored or created wetlands have shown significant 
increases in numbers of obligate wetland species, num- 
bers of individuals, and rare obligate wetland species 
(Dick 1993; Hickman 1994; Pollard et al. 2000). Of 
the rare birds observed at Hilliardton Marsh, of par- 
ticular importance are those that are provincially sig- 
nificant. American Coot, Black Tern, and Yellow Rail 
are rare breeders in northern Ontario (Cadman et al. 
1987). Amerian Coot and Black Tern have declined in 
numbers and distribution over the past two decades 
(Cadman et al. 1987; Austen et al. 1994), and Yellow 
Rails are among the most reclusive birds in Ontario 
(Bart et al. 1984). 

It is also significant that observations and abun- 
dances of many species of rare birds did not change 
between years. Open, riparian habitat for the faculta- 
tive wetland LeConte’s Sparrow (Cadman et al. 1987) 
lost during wetland construction was recreated during 
the same process in different locations. Open, prairie- 
type habitat that is suitable for Horned Lark and 
Eastern Meadowlark (Cadman et al. 1987) was main- 
tained. 

Additionally, some new and rare birds, mostly at the 
local level, were observed from the facultative, fac- 
ultative upland, and upland groups during 1998. These 
included Warbling Vireo, Gray Catbird, Baltimore 
Oriole, and Brown Thrasher (Table 3). Some rare spe- 
cies such as Upland Sandpiper and Connecticut Warb- 


Locky, DAVIES, AND WARNER: EFFECTS OF WETLAND CREATION BB 


@@M—™ Obligate Wetland 
SSSS Facultative Wetland 
@@@A Facultative 
Facultative Upland 
@@m@mm@ Upland 


Species Richness 


Moderate 


High 


No Change 
Degree of Habitat Change 


FIGURE 3. Degree of habitat change after wetland construc- 
tion and bird species richness by bird groups with 
95% confidence interval error bars. See text for 
explanation of bird groups. 


ler observed in 1993 were not observed in any of the 
post-construction surveys, and this could be attribut- 
able to natural variation within the bird population 
(Holyoak and Baillie 1996). We were reasonably con- 
fident that varying weather conditions; e.g., drought, 
was not a factor affecting bird species abundance be- 
tween years. Long-distance migrants such as Oven- 
birds and Red-eyed Vireos are often more susceptible 
to drought (Blake et. al. 1992); numbers recorded in 
the control stations were stable between years and 
precipitation records for the study period indicated 
no anomalies. 

Continued monitoring of the bird community at 
Hilliardton Marsh would be necessary to determine 
long-term trends at the species level, especially for 
rare species. Additionally, new wetland projects run 
the risk of eradication of existing important habitat 
or have negative impacts on rare species (Keddy and 
Wisheu 1989; Hickman 1994). Maintaining habitat 
amenable to all rare species should be a high man- 
agement priority at Hilliardton Marsh. 


Conclusions 

This study provides an initial view of the response 
of the bird community during the breeding season to 
wetland construction in a region where there have 
been few studies on birds. Bird diversity increased sig- 
nificantly in response to wetland construction, espe- 
cially wetland birds. Constructed wetlands are not 
always successful in augmenting the bird community 
and the intended use and consequences must be care- 
fully planned (Zhijun et al. 2004). The original aim 
of creating Hilliardton Marsh was to provide water- 
fowl habitat, and this and related work (Locky 1999) 
have shown this aim to be met. However, wetland con- 
struction has also attracted a range of non-target, non- 
waterfowl wetland species, while maintaining the 


74 THE CANADIAN FIELD-NATURALIST 


natural diversity of the original upland bird commu- 
nity, including rare species. A number of new wetland 
rare species were also attracted. This construction has 
been particularly favorable because the marsh habi- 
tat that was introduced is rare in the region. There- 
fore, the type of habitat being created is an important 
consideration when designing wetland construction 
projects and is also a consideration for continued man- 
agement of constructed wetland systems. Like all 
wetlands, constructed wetlands are dynamic systems 
and long-term management and surveys would be 
required to ensure that the enduring potential of pro- 
viding wetland bird habitat is maintained. 


Acknowledgments 

Alison Buchanan and Bruce Murphy assisted in 
the field and Doug Brook and Peter Davis provided 
helpful discussion. Valuable comments provided by 
A. J. Erskine, S. R. Wilkinson, M. N. Thormann, S. 
J. Hannon, S. E. Bayley, and an anonymous reviewer 
substantially improved this manuscript, and were 
greatly appreciated. This study was supported by the 
Northeast Science and Technology Unit of the Min- 
istry of Natural Resources, Ducks Unlimited Canada, 
and the Natural Sciences and Engineering Research 
Council of Canada. 


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Temiskaming Field Naturalists. 1994. Birds of the Little 
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Received 15 March 2004 
Accepted 7 February 2005 


Lichen Trimlines in the Peace-Athabasca Delta: Variations in Flora, 
Form, and Disturbance Regime 


KEVIN P. Timoney! and JANET MARSH? 


'Treeline Ecological Research, 21551 Twp Road 520, Sherwood Park, Alberta, T8E 1E3 Canada; e-mail: ktimoney@compu 
smart.ab.ca 
2040 7" Avenue NE, Calgary, Alberta, T2E ON8 Canada; e-mail: Marshj4@aol.com 


Timoney, Kevin P., and Janet Marsh. 2005. Lichen trimlines in the Peace-Athabasca Delta: variations in flora, form, and dis- 
turbance regime. Canadian Field-Naturalist 119(1): 76-81. 


Lichen trimlines are characteristic of aquatic systems where lichen-covered rocks border fluctuating water bodies. This study 
examined water-origin saxicolous lichen trimlines on acidic metacrystalline bedrock outcrops in the Peace-Athabasca Delta, 
northern Alberta, Canada. Twenty-seven species of saxicolous lichens were found in the vicinity of the trimlines. Species 
richness above the trimline (26 species) was almost twice that found below the trimline (14 species). Colonization lag time, 
differences in susceptibility to disturbance, and site influences on lichen establishment and survival might be involved in the 
absence of many species below trimlines. In frequently inundated areas, rock surfaces are dominated by the amphibious lichens 
Staurothele fissa and S. drummondi. The dominant lichen colonizer below trimlines was PAyscia caesia. Other important 
colonizers below trimlines included Phaeophyscia sciastra and Physcta dubia, and on rocks fertilized by bird feces, Xanthoria 
elegans. A curious feature of the saxicolous flora was the presence of many calciphiles. Variations in trimline form and height 
and constituent species are related to the hydrologic and disturbance regime. Trimlines in perched basins tend to be near 
current water level, horizontal, and distinct, and indicate a relatively stable, infrequently flooded environment. Trimlines at 
open-drainage sites tend to be high above current water, wavy, and indistinct, and indicate a strongly-pulsed environment 


with frequent disturbances. Those at restricted-drainage basins are variable in form and height and disturbance regime. 


Key Words: delta, disturbance, Peace-Athabasca, saxicols, trimline, water regime, wetland, zonation, Alberta. 


Zonation of lichen communities along lake and river 
shores has been noted for many decades (e.g., Santes- 
son 1939) and has been attributed to gradients in phy- 
sical and ecological factors. Lichen species differ in 
their tolerance of immersion/desiccation, siltation, 
water and substrate chemistry, water and ice scouring, 
current velocity, substrate stability, wave splash, shade, 
competition with bryophytes and vascular plants, and 
colonization rates after disturbance (Scott 1967; Ried 
1974; Rosentreter 1984; Beckelhimer and Weaks 1984, 
1986; Pereira et al. 1987; Gilbert and Giavarini 1997, 
2000). While naming conventions have differed, lich- 
enologists have consistently noted four lichen zones 
along lakes and rivers; e.g., for British lakes, Gilbert 
and Giavarini (2000) used the zones “‘terrestrial’’, “up- 
per splash’, “lower splash”, and “submerged”, while 
for the lower reaches of the Salmon River, Idaho, USA, 
Rosentreter (1984) used the zones “extreme flood”, 
“high flood”, “normal flood”, and “low water”. 

Saxicolous lichen trimlines are typically distinct 
transition zones on bedrock-lined shores, below which 
lichens intolerant of immersion may be scarce or ab- 
sent (Timoney and Marsh 2004). Above the line, there 
may be a continuous lichen flora attached to the rock 
surface; below the line there may be dominance by 
amphibious lichens such as Verrucaria or Staurothele 
(Brodo et al. 2001). In terms of typical lichen zonation, 
a trimline would be located somewhere in the “upper 
splash” / “high flood” zone. 


76 


Worldwide, there are two fundamentally different 
types of lichen trimlines: those due to scouring by 
glacial ice and those due to aquatic disturbances (e.g., 
Winchester and Harrison 2000). The latter are typi- 
cally due to flooding or sedimentation that results in 
lichen mortality, followed by a low water period that 
exposes the transition between living lichens and bare 
rock or tree bark (Hale 1984; Beckelhimer and Weaks 
1984, 1986). 

Saxicolous lichen trimlines are characteristic of 
aquatic systems where lichen-covered rocks border 
fluctuating water bodies, e.g., in Florida (Hale 1984), 
in the Geraldine Lakes area of Jasper National Park, 


Alberta, and at various lakes in the Boundary Waters | 
Canoe Area, Minnesota, USA (Timoney, field obser- — 


vations), in New South Wales, Australia (Gregory 
1976), and on Georgian Bay, Ontario (Mitchell 2002). 

In this era of generally declining water levels in 
much of North America, lichen trimlines may be use- 
ful as datable markers of previous high water levels. 
The objectives of this note are to document the lichen 
species growing on rock near lichen trimlines in a 
strongly-pulsed northern wetland and to describe vari- 
ations in trimline form in the context of ecological 
and hydrological factors. 


Study Area 


The study area is located in the northeast portion 


of the Peace-Athabasca Delta, northern Alberta. While | 


ee 


2005 


the delta is located within the Continental High Boreal 
wetland region of Canada (National Wetlands Work- 
ing Group 1988), its biota and vegetation are charac- 
teristic of the Continental Prairie wetland region 
(Timoney 2002). Inclusive of Point Providence and 
the Scow Channel areas, the total area of the delta is 
5168 km?, composed of 2406 km? of water and 2762 
km? of non-water. The non-water portion of the delta 
includes forests, shrublands, marshes, meadows, mud- 
flats, peatlands, and bedrock knolls. The present, dom- 
inant non-aquatic vegetation of the delta includes 
Carex atherodes (Awned Sedge) and Scoloch/oa Jes- 
tucacea (Spangletop) marshes, Ca/amagrostis canaden- 
sis (Bluejoint Reedgrass) meadows, Sa/ir (Willow) 
thickets and savannahs, and forests of Populus bal- 
samifera (Balsam Poplar) and Picea glauca (White 
Spruce) (Timoney 2004*). 

Drainage basins are classified as hydrologically 
“open” (connected to Lake Athabasca or to major rivers 
at all times); semi-restricted (connected only at times 
of high water); or perched (receiving water only during 
ice-jam floods, through local precipitation or by local 
drainage within their catchments). Bedrock outcrops 
border many of these basins and are composed pri- 
marily of Precambrian acidic metacrystalline rocks 
(granitic or felsic gneisses). Many of these bedrock 
outcrops exhibit saxicolous lichen trimlines. 


Methods 

Sixteen sites that represented three basin types 
(open, semi-restricted, and perched) were sampled 
from 17-21 June 2002 and from 25-29 August 2003 
(Table 1, Figure 1) as part of a reconstruction of water 
levels by lichenometry (Marsh and Timoney 2003). 
Access to the sites was by boat or helicopter (Site 5 is 
accessible on foot). Each site consisted of a boulder 
outcrop that exhibited a lichen trimline. At each site, 


TABLE |. Location (decimalized latitude and longitude) and 
basin type of the trimline study sites. 


Site Latitude Longitude Basin Type 
] 58.497 719.593 open 
2 58.546 119.605 perched 
3 58.532 119.601 semi-restricted 
4 58.342 119.555 semi-restricted 
> 58.420 119.574 open 
6 58.408 Bis Pome | open 
7 58.497 119.593 open 
8 58.522 119.599 perched 
“] 58.558 119.608 perched 
10 58.577 119.613 semi-restricted 
11 58.610 119.621 perched 
12 58.608 119.620 perched 
13 58.666 119.635 perched 
14 58.658 119.633 semi-restricted 
15 58.518 119.598 perched 
4 58.521 119.599 semi-restricted 


 ——— CO eee 


TIMONEY AND MARSH: LICHEN TRIMLINES IN THE PEACE-ATHABASCA DELTA 5 A | 


FIGURE |. Top: Lichen study sites in the northeast sector of 
the Peace-Athabasca Delta (geographic center at 
~111°20'W, 58°48'N). The asterisk * between Sites 
3, 8 and 15 refers to “lichen observation site”. Width 
of image from east to west is ~ 66 km. Bottom: 
regional context of study area. 


a bench mark (a galvanized nail with orange flagging) 
was placed on the trimline in the middle of the area that 
was to be surveyed. The site was plotted on an NTS 
1:50000 map, from which UTM coordinates were 
derived. The vertical height of the lichen trimline was 
measured with a clinometer (used as a level) sighted 
on a tape measure at the water’s edge. Lichen species 
present near the trimline were recorded or collected 
for later identification. Sampling focused on foliose 
and crustose species attached to rock below the trim- 
line and within one vertical meter above the trimline 
(i.e., roughly, in the “upper splash” / “high flood” and 
the “terrestrial” / “extreme flood” zones). For each spe- 
cies, </=15 lichen thalli diameters were measured to 
the nearest mm. The choice of thalli was not random; 
we looked for the larger thalli. Site descriptions of 
trimline aspect, width, elevation, slope, and vegeta- 
tion community were prepared and photographs were 
taken. We described trimline form and noted relevant 
site and disturbance factors. Lichen nomenclature fol- 
lows Esslinger (1997*), and that for vascular plants 
follows Moss (1983). 


Results 
Flora 

The saxicolous lichen flora included at least 27 spe- 
cies (Table 2). Twenty-six species were found above the 
trimline and 14 species were found below the trimline. 
Only one species, Staurothele drummondi, appeared to 
be restricted to the zone below the trimline although 
Staurothele fissa was more abundant below the trim- 
line than above it. There was a strong compositional 
break at the trimline. Many species common above 
trimlines were not found below trimlines, e.g., Aspic- 
tla, Dimelaena, Dermatocarpon, Lasaltia, and Umbi- 
licaria and some species of Melanelia and Parmelta. 
Other species found above and below the trimline were 
far more abundant above the trimline (e.g, Xavthopar- 
melita somlo€nsis). 

Below the trimline, rock surfaces were usually dom- 
inated by Staurothele drummondi or S. fissa, dark 
brown, thin crustose lichens with or without an assem- 
blage of colonizing lichens such as Physcia caesta. At 
the majority of outcrops, the lichen community within 
circa one meter above the lichen trimline (the dry zone) 
was dominated by the foliose lichens Xanthoparmelia 
somloénsis, Physcia caesta, P. phaea, Phaeophyscia 
sclastra, Melanelia stygia, Rhizoplaca chrysoleuca, 
Umbilicaria deusta and the crustose lichens R/zzo- 
carpon disporum, Dimelaena oreina and Aspicilia cae- 
stocinerea. Beyond the reach of extreme high water, 
were a variety of lichen communities, e.g., fruticose 
mat formers (e.g., many Cladonia/Cladina species) or 
Umbilicarta-dominated areas, or vascular plant and 
moss communities (e.g., /wiperus (Juniper), Saxi/raga 
tricuspidata (Prickly Saxifrage), Amelanchier alnifo- 
fia (Saskatoon), Agvopyron trachycaulum (Slender 
Wheatgrass), Polyirichum, Drepanocladus, etc.) that 
lay beyond the scope of this study. 


Variations in Trimline Form, Disturbance Regime, 
and Flora 

Typical trimline form in open basins was wavy and 
indistinct; that in perched basins was level and distinct; 
in semi-restricted basins trimline form was variable 
(Table 3). Median trimline heights in open and semi- 
restricted basins were >2 m above current water levels, 
while perched basin trimline heights were <1 m above 
basin water levels. Median trimline elevations were 
similar across types while water elevations in open and 
semi-restricted basins were lower than in perched basins. 

We observed seven variations in the trimlines. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE 2. Saxicolous lichen species observed at the 16 study 
sites. 


Above 
Trimline 


Below 


Species Trimline 


Aspicilia caestocinerea 
Cladina stellaris 
Dermatocarpon reticulatum 
Dimelaena oreina 

Lasallia pensylvanica 
Leproloma vouauxit 
Melanelia disjuncta 
Melanelia sorediata 
Melanelia stygia 

Parmelia saxatilis 
Parmelia sulcata 
Phaeophyscia sciastra 
Phaeophyscia hispidula 
Physcia caesta 

Physcia dubia 

Physcia phaea 

Physconia muscigena 
Placidium squamulosum 
Ramatlina intermedia 
Rhizocarpon disporum 
Rhizoplaca chrysoleuca 
Staurothele drummondii 
Staurothele fissa 
Umbilicaria deusta 
Umbilicaria muehlenbergii 
Xanthoparmelia somlo€nsis 
Xanthoria elegans 


* 


CoCo el ait YC Yas YOM TOS flea Gc HG VEC Tks Om SY Pa EK WR Tt 


Ss = SS 


At Perched and Semi-Restricted Basins: 

1. In most perched basins, trimlines were level and distinct 
with a characteristic flora of colonizing lichens below the 
trimline indicative of infrequent water level fluctuations 
(Figure 2a). 

2. In two perched and semi-restricted basins, there were no 
colonizing lichens below a trimline located near the cur- 


rent standing water. The below trimline communities were | 
dominated by amphibious Svauwrothele fissa and/or S. drum- | 


mondit. Such sites indicate frequent small water level 


fluctuations. At Site 10 (Figure 2b), water levels were influ- — 


enced by beavers. Dominance by lichens that favor periodic 
submergence and the absence of other lichens argues for 
periodic inundation. At Site 12, a high elevation, isolated 
perched basin (Figure 2c), the lowermost S‘aurvorhele was 
submerged 20 cm at time of sampling (19 June 2002). 

3. At Site 11 (Figure 2d), the mid-lake outcrop served as a 
resting area for ducks and black terns (Ch/idonias niger). 
Nitrophilic Xanthoria elegans and amphibious Staurothele 
were the only lichen species present below the trimline. 


TABLE 3. Trimline form, height, elevation and water elevation by basin type. 


Trimline Ht Trimline Elevation Water Elevation 
Basin Type (n) Trimline Form (median, m)* (median, m asl) (median, m asl) 
open (4) wavy, indistinct 2.20 210.83 208.61 
semi-restricted (5) variable 2518 210.44 208.29 
perched (7) level, distinct 0.91 210.78 209.84 


* above water level at time of sampling 


ee 


2005 TIMONEY AND MARSH: LICHEN TRIMLINES IN THE PEACE-ATHABASCA DELTA 719 


FIGURE 2(a) A distinct and level trimline at perched basin Site 2, 79 cm above water at time of sampling. P/yscia caesia was only col- 
onizing lichen below the trimline. (b) A Stawrothe/e community at a beaver-dam influenced semi-restricted basin (Site 10) with 
a trimline 40 cm above water. (c) At this isolated, bedrock upland perched basin (Site 12), above the trimline (left half of photo), 
dominants are Rhizocarpon, Xanthoparmelia, Aspicilia, and Me/ane/ia, At the indistinct trimline (60 cm above water), there 
is a lichen-free whitish band, followed by a blackish Staurothele band, and below that, extending under water, a mixture of 
oxide-staining and S/aurorhe/e thalli. (d) A level trimline influenced by bird feces at perched basin Site 11, 102 cm above 
water. Above the trimline (white line) Xanthoria elegans dominates the cover (its orange thalli do not contrast well against 
the bedrock in the gray scale image); Physcra caesia is the only other colonizing lichen present. The dark band below the 
trimline is composed of Szaurothe/e. (e) A high, indistinct, wavy trimline (white line near the backpack) at Site 5 on Lake 
Athabasca, 224 cm above water. Dominant colonizing lichen below the trimline was PAyscia caesia, accompanied by amphib- 
ious S7aurothe/e (white S) near water level. (f) A wavy and indistinct trimline at Site 7, 220 cm above water, along the Rochers 
River. Note the patch of flaked rock (pointer from P) in which several young lichens have established. (g) A Leproloma vouauxti 
trimline on a moist and shady vertical to overhanging gneiss at Site 4, behind a levee, 230 cm above river level. (h) A high, 
distinct level trimline at the semi-restricted “*” site, 218 cm above water. Note the band of lichen-free white weathered rock 
below the trimline, below that the blackish S7awrothe/e band, and the lowest band (light gray, but reddish orange in color) of 
iron oxide stained rock. 


80 


At Open (and Semi-restricted) Basins and Riparian 

Sites: 

4. Open basins were typified by high and often indistinct and 
wavy trimlines (Figure 2e). Colonizing lichens were present 
or absent, but Sauvorhe/e was present near the current water 
level. The high trimlines indicated a large drawdown, while 
the indistinct and wavy trimlines indicated episodic inun- 
dation or disturbance by e.g., open water and ice-jam floods, 
seiches, and ice scour. The disturbance regime at Lake Atha- 
basca sites would be influenced by aspect and exposure to 
prevailing winds, waves, and ice scour. 

5. The typical active riparian trimline was high and sometimes 
indistinct and wavy, indicative of active disturbance. At 
Site 7 on the Rochers River, patches of unweathered, lichen- 
free rock existed alongside rock covered by large, mature 
lichens and patches with young, newly-establishing lichens 
(Figure 2f). Colonizing lichens and S7aurothele spp. may 
be present or absent at such sites. 

6. Site 4 (Figure 2g), along a semi-active branch of the Quatre 
Fourches River not subject to wave or ice action, was a 
moist, shady seepage site of vertical to overhanging beds 
of gneissic bedrock. There was nearly complete cover of 
Leproloma vouauxti above the trimline and a sparse cover 
below the trimline; Ramalina intermedia and Physcia caesta 
were present above the trimline. 

7. At two semi-restricted sites, we found no colonizing lichens 
below a level, high, distinct trimline on weathered granite, 
below which only S*aurorhele grew (Figure 2h). The ab- 
sence of non-amphibious lichens indicates an unexplained 
impediment to lichen colonization and/or survival. The area 
is subject to seiches; e.g., from evening 18 June to evening 
19 June 2002, local water level fell 30 cm. But seiches of 
a 2 m magnitude would be difficult to envision and would 
not be of sufficient duration to kill lichens. 


Discussion 

While the local bedrock is nominally acidic, near the 
trimlines, many lichens, mosses, and vascular plants 
indicate basic or circumneutral soil conditions. Some 
examples are, amongst the lichens — Xanthoria ele- 
gans, Physcia caesta, Dermatocarpon reticulatum, and 
Placidium squamulosum (Vitt et al. 1988); and mosses 
— Thuidium abietinum, Tortula ruralis, Rhytidium 
rugosum, and Encalypra sp. (Ireland 1982). Amongst 
the neutral to basic soil indicators in the vascular plants, 
Picea glauca dominates the forests near the trimlines. 

There is no clear reason for the commonness of cal- 
ciphiles on nominally acidic bedrock. We observed 
“dust” on rocks and saxicolous plants in the area which 
may be calcareous loess deposited on the outcrops 
rocks when nearby mudflats dry and are exposed to 
wind. In other situations, wave splash and wind-driven 
spray may wet the surfaces with delta water. Delta 
surface water is nutrient-rich and alkaline (Hall et al. 
2003*; e.g., mean pH of lakes = 8.2, of rivers = 8.0). 
We found no data on the pH of local rainfall. 

At many sites, the most abundant lichen below the 
trimline was Physcia caesia (a nitrophile; McCune and 
Geiser 1997; Brodo et al. 2001). Physcia caesia, 
Xanthoparmelia somloénsis, and Staurothele fissa were 
common pioneer species in both perched basins and 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


open sites whereas Phaeophyscia sciastra colonized 
only perched basins. Primary sites for colonization were 
cracks; seepage, rough, and protected shady/moist sur- 
faces; and dark-colored mineral bands. As birds often 
rest on shore-side rocks, nitrogenous bird waste may 
play a role in both colonization and species composi- 
tion of the saxicolous communities. 

Wide variations in water level, and ice-scour, along 
the rivers and Lake Athabasca may lead to the indis- 
tinct and wavy trimlines characteristic of open and 
some semi-restricted sites. At many sites, below the 
trimline there was a light-colored rock band situated 
above a reddish-orange band of oxidized iron (e.g., 
Figure 2c, h). This color pattern may indicate a tran- 
sition from frequent reduction and oxidation due to 
water level fluctuations (reddish-orange) to infrequent 
inundation (light colored rock). Ice scour by wind- or 
river-driven ice blocks may scrape the rocks free of 
lichens, especially foliose species that are not appressed 
to the rock. The chaotic piling of ice blocks character- 
istic of break-up might contribute to the irregularity of 
the trimlines on open drainage sites. Flaking of the bed- 
rock, perhaps by ice-blocks pulling away from the rock, 
or by 77 situ formation of ice lenses, might also act to 
delay lichen succession. 

Width of the trimline, slope, aspect, and basin type 
were not related to the number of species found below 
the trimline (Marsh and Timoney 2003*). While site 
factors such as aspect can be important in determining 
the distribution and performance of lichens (Beckel- 
himer and Weaks 1986; Eversman 1982), both the rel- 
atively short time for establishment and the humidity 
along the shores may have masked physical effects. 

Gregory (1976) found double trimlines along a river 
in Australia. We found no double trimlines (indicative 
of a high flood, followed by a period of lichen estab- 
lishment below the trimline, followed by a lower flood 
that results in a second, lower trimline). Fourteen of 


16 trimline elevations fell within a narrow range of | 
210.26-210.92 m asl. Based on lichen thalli diameters _ 


and local growth rates, Timoney and Marsh (2004) 
surmised that most of the trimlines probably formed 
during the period 1976 to 1990. Many of the trimlines 
probably formed during relatively-persistent high 
water which followed a major 1974 ice-jam flood. The 
drought-prone early 1980s may have been a key time 
of lichen establishment below trimlines (Marsh and 
Timoney 2003*). The lichen data bear on the contro- 
versy surrounding the history of changing water levels 
in the Peace-Athabasca Delta. As the trimlines estab- 
lished in response to flooding that occurred after the 


a 


—— 


ee 


construction of the Bennett Dam, the trimlines can- | 


not be construed as pre-dam high water marks. The | 
reverse is true—they are post-dam high water marks. | 

How long might trimlines persist in the absence of | 
a subsequent rise in water level that would obliterate | 
colonizing lichens? Thalli below trimlines might reach _ 


a mean diameter within one standard deviation of © 


2005 


above-trimline diameters in about 27 to 55 years of 
uninterrupted growth. Allowing for a lichen establish- 
ment lag time of about 6-10 years in the delta (Marsh 
and Timoney 2003*), trimlines there might remain 
visually distinct for roughly 33-65 years. 

Some sites had no datable lichens (only Saurothe/e) 
below the trimline. Species differences in colonization 
rates (Gilbert and Giavarini 2000) of these relatively 
recent surfaces, differences in susceptibility to distur- 
bance, and site influences on lichen establishment and 
survival might be involved in the absence of many spe- 
cies below trimlines. As sites that offer insight into 
lichen colonization, succession, lichenometry, water 
level variations, and disturbance ecology, lichen trim- 
lines deserve more study. 


Acknowledgments 

Thanks to Irwin Brodo, James Case, Alan Fryday, 
D. J. Hill, Derek Johnson, Anne Robinson, Roger Ros- 
entreter, Mark Seaward, and Pat Wolseley for their 
comments regarding lichen zonation/trimlines, lichen- 
ometry, and lichen susceptibility to immersion. BC 
Hydro and Wood Buffalo National Park Fort Chipew- 
yan staff provided logistic support. Stephen Hamilton 
assisted with office tasks. 


Documents Cited (marked * in text) 

Esslinger, T. L. 1997. A cumulative checklist for the lichen- 
forming, lichenicolous and allied fungi of the continental 
United States and Canada [online]. North Dakota State 
University. Available from: http://www.ndsu.nodak.edu/ 
instruct/esslinge/chcklst/chcklst7.htm. [updated 17 July 
2002; cited 4 February 2004] 

Hall, R., B. Wolfe, and T. Edwards. 2003. A multi-century 
flood, climatic, and ecological history of the Peace-Atha- 
basca Delta, northern Alberta, Canada. Final report to BC 
Hydro, Burnaby, British Columbia. 

Marsh, J., and K. Timoney. 2003. Saxicolous lichen trim- 
lines in the Peace-Athabasca Delta of Northern Alberta: 
flora, growth rates, establishment and persistence, and his- 
tory of water levels. Final report to BC Hydro, Burnaby, 
British Columbia. 

Timoney, K. P. 2004. Vegetation, climate, and biophysical 
variation and change in the Peace-Athabasca Delta, 1993- 
2003. Final report to BC Hydro, Burnaby, British Columbia. 


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, The Lichenologist 18: 339-347. 

| Brodo, I. M., S. D. Sharnoff, and S. Sharnoff. 2001. 

_| Lichens of North America. Yale University Press, New 

Haven, Connecticut. 


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Gilbert, O. L., and V. J. Giavarini. 1997. The lichen vege- 
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(Granada, S. de Espana). Cryptogamie, Bryologie, et Lich- 
enologie 8: 263-273. 

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Rosentreter, R. 1984. The zonation of mosses and lichens 
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108-117. 

Santesson, R. 1939. Uber die Zonationsverhiltnisse der 
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Timoney, K. P. 2002. A dying delta? A case study of a wet- 
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Timoney, K. P., and J. E. Marsh. 2004. Lichen trimlines in 
northern Alberta: establishment, growth rates, and historic 
water levels. The Bryologist 107: 429-440. 

Vitt, D. H., J. E. Marsh, and R. B. Bovey. 1988. Mosses, 
lichens & ferns of northwest North America. Lone Pine 
Publishing. Edmonton, Alberta. 

Winchester, V., and S. Harrison. 2000. Dendrochronology 
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Patagonia Icefield, Chile. Geomorphology 34: 181-194. 


Received 12 May 2004 
Accepted 4 March 2005 


Identification of a Marine Green Alga Fercursaria percursa from 
Hypersaline Springs in the Middle of the North American Continent 


KATHLEEN L. Lonpry!, PASCAL H. BADIoU2, and STEPHEN E. GRASBY? 


'Department of Microbiology, University of Manitoba, Winnipeg, Manitoba, R3T 2N2 Canada; e-mail: londryk @cc.umanitoba.ca 
"Department of Botany, University of Manitoba, Winnipeg. Manitoba, R3T 2N2 Canada; e-mail: pbadiou@shaw.ca 
3Geological Survey of Canada, 3303 33™ St. NW, Calgary, Alberta, T2L 2A7 Canada; e-mail: SGrasby@NRCan.gc.ca 


Londry, Kathleen L., Pascal H. Badiou, and Stephen E. Grasby. 2005. Identification of a marine green alga Percursaria percursa 
from hypersaline springs in the middle of the North American continent. Canadian Field-Naturalist 119(1): 82-87. 


The chlorophycean alga Percursaria percursa (Ulvaceae, Ulvales, Chlorophyceae), typical of marine inter-tidal zones, is reported 
for the first time from hypersaline springs located along the north-western shore of Lake Winnipegosis in Manitoba. Although 
not usually found inland, P yercursa is the dominant member of microbial mat communities that thrive in shallow pools at the 


outlets of hypersaline springs. 


Key Words: Percursaria percursa, chiorophyta, hypersaline, springs, marine algae, Manitoba. 


This study was undertaken to characterise the mem- 
bers of microbial mat communities growing in the 
pools and outflow channels of hypersaline springs in 
North-Western Manitoba. Studies on the microbial 
ecology of hypersaline waters has previously been lim- 
ited to marshes and standing bodies of water such as 
salt lakes and salt production ponds (Oren 2002), so 
very little is known about the microbes inhabiting flow- 
ing water sources such as springs. Stable hypersaline 
springs are uncommon, and well-defined springs amen- 
able to scientific study, particularly at low temperatures, 
are relatively rare. However, these springs can provide 
important information about the survival of microbes 
under extreme conditions. They also provide a modern 
analogue for the extensive microbial mats that charac- 
terised life on Earth during the Precambrian. 

The highly saline pools, marshes, and saltpans on 
the northern margins of Lake Winnipegosis, Manitoba, 
have created a unique ecosystem dominated by organ- 
isms characteristic of marine habitats (Patterson et al. 
1997). The flora and fauna have been described pre- 
viously, with plant species composition similar to 
prairie saltpans (Burchill and Kenkel 1991), and ani- 
mals that included marine representatives (Chordata, 
Protozoa, Rotifera, and Arthropoda) (McKillop et al. 
1992). The distribution of the vegetation was linked to 
the salt tolerance of the species (Burchill and Kenkel 
1991). It has been suggested (Patterson et al. 1997) 
that these marine species were transported to the springs 
area in Manitoba by birds (including Canada Geese) 
from coastal areas such as Hudson Bay, 700 km north- 
east of this site, and the Gulf of Mexico. Coloniza- 
tion of the saline springs could be no sooner than the 
retreat of the continental ice sheet, which covered the 
region up until about 10 000 years ago. Patterson et 
al. (1997) show that a saline ecosystem has been in 
place since at least 5500 years ago. Colonization by 


marine species occurred soon thereafter (Patterson et 
all 1997). 

Geochemical and stable-isotope analyses indicate 
that spring waters originate as meteoric water (Gras- 
by et al. 2000). There was a large influx of glacial melt- 
water into the underlying bedrock during the Pleis- 
tocene, which dissolved the underlying beds of halite 
(NaCl), sylvite (KCI) and similar salts deposited by 
evaporation of Devonian seawater. The retreat of ice 
sheets has released the waters, discharging now as | 
saline springs (Grasby et al. 2000; Grasby and Chen | 
2005). There are numerous saline springs along the 
western shore of Lake Winnipegosis (Grasby 2000). | 
Microbial mats have only been observed at springs in | 
the Dawson Bay area (on the north-western shore of 
the lake), further west along the Red Deer River (Mani- | 
toba), as well as springs near Pelican Bay (east of Daw- — 
son Bay). These hypersaline springs flow over a saltpan | 


of barren, iron stained surficial material (McKillop et | 


al. 1992). The high salt concentrations severely limit 
plant growth in the area, and life within the springs — 
themselves is restricted to microorganisms. 


Study Site ! 
The hypersaline springs are located along the west- | 
ern shore of Lake Winnipegosis in North-Western | 
Manitoba. As shown in Figure 1, the springs are in | 
Manitoba, just south and west of Dawson Bay in the | 
northern extent of Lake Winnipegosis. The locations | 
and chemistry of the spring outlets are summarised in | 
Table 1. Water samples were collected and preserved | 
in the field. The water was passed through a 0.45 um | 
filter. Samples for Fe analysis were acidified to pH. 
<2 with ultrapure NHO,. Samples for anion analysis | 
were untreated. Anions were measured by ion liquid. 
chromatography, and Fe by ICP-MS. The salinity of | 
springs studied here, the feature that defines the springs, 
| 


2005 


na 
2 SASKATCHEWAN & 


32] 
N 


51° 


50°} 


4:3,000,000 
49° 


100° 99° 98° oF 


LONDRY, BADIOU, AND GRASBY: IDENTIFICATION OF A MARINE GREEN ALGA 83 


ONTARIO 


@ saline spring 


O spring with extensive 
mat development 


¥& spring with Percursaria 
percursa confirmed 


FiGurE |. Regional map showing locations of saline springs, indicating which ones have been observed to have microbial 


mats of 2 percursa. 


and limits growth in these areas, ranges from below 

seawater to twice seawater concentrations. The salinity 

is mostly from NaCl (from halite), with minor differ- 
ences in composition compared to typical seawater. 

The salt from these springs has been used historically 

by early settlers (Petch 1987). Chloride and sulfate 

concentrations in springs containing microbial mats 
are typically 24 to 36 g/l and 2.1 to 3.5 g/l, respectively 
(Table 1). Gas bubbles rising from the spring sediments 
were observed frequently, often becoming trapped by 
the overlying mat in springs with confluent growth, 
leading to the appearance of medium to large swellings 
of the mat fabric. The gas composition is dominantly 
N, (95%) with traces of CO, (1.8%), He (1.5%), O, 
(1.0%), Ar (0.8%), and CH, (0.02%) (McKillop et al. 
1992), with the latter two potentially supporting micro- 
- bial growth. 

Each site consisted of multiple spring outlets, most 
of which formed pools around the origin, the oldest of 
| which have formed raised cauldrons. Microbial mats 
| grew on the surface and around the margins of the 
pools (Figure 2). There was considerable red goethite 
| (FeO*OH) staining in areas surrounding the springs, 
| but not in the spring sources themselves, which are 
|| dominated by calcite and quartz. Areas surrounding 
the springs also sometimes exhibited a characteristic 
“brainy” texture due to the decomposition of the algal 
mats beneath the evaporitic gypsum crust. The springs 


flow year-round and the salinity keeps some of the 
larger ones from freezing, even in winter (McKillops 
et al. 1992). Microbial mat growth was observed in 
freezing temperatures in the fall and spring. Samples 
of the microbial mats were collected in October 2001, 
May 2002, and August 2003. Samples were stored on 
ice or in a freezer (-20°C) until they were analysed 
by microscopy at the University of Manitoba using a 
Leitz Diaplan (Germany) microscope. 


Results and Discussion 
Organism 

Percursaria percursa (C. Agardh) Bory De St. Vin- 
cent is a green alga of the order Ulvales (Ulvophyceae, 
Chlorophyta) that is phylogenetically related to U/va 
intestinalis (O’ Kelly et al. 2004). It forms distinct 
biseriate simple threads of varying breadth (Bliding 
1963). The thalli are typically unbranched, up to 3 cm 
long, flexuous and contorted. The double row of cells 
is usually in exact symmetry (Figure 3), with each 
cell containing a single parietal band-like chloroplast 
similar to those seen in species of the genus U//orhrix. 
Under intense light, the chloroplast forms a narrow 
parietal girdle in the middle of the cell, whereas under 
low light levels the chloroplast becomes well devel- 
oped and less band-like (Bliding 1963). The number 
of pyrenoids in the chloroplasts of P percursa varies 
between one and three with most specimens typically 


84 


TABLE |. Locations and physicochemistry of spring waters west of Lake Winnipegosis. 


THE CANADIAN FIELD-NATURALIST 


sample Latitude Longitude 
number NAD 87 

M993013? 52°48.105' 100°48.750' 
M003005 52°47.977 100°49.430' 
M99303 1 52°40.893' 100°4.798' 
M993016? 52°45.086' 100°52.926' 
M993029? 52382235. 100°22.120' 
M993028? 52°41.561' 100°21.176' 
M993027? 32, 41.259 100°20.988' 
M993019° Seok is2 101°08.166' 
M993014° 52°47.703' 100°51.817' 
M003003 * 2 103" ~ 101°04.169' 
M993020° 52, 209 101°08.159' 
MO003006° 52°47.919' 100°53.118' 
M993026? 32°91. 783" 101°05.392' 
M983075 ? 3252/0352 101°03.271' 
M993011° 52°52.603' 101°02.886' 
M993025 52°45.806' 100°52.845' 
M003002* 52752191" 101°05.563' 
M993037 51°37 A429! 99°58.472' 
M983076? 52°92.926' 1OTO 22359 
MO003004 ° 52°45.266' 100°53.429' 
M993038 51'°491099' 99°43.493' 
M993035 51°37.424' 99°56.777 
M993034 51°43.390' 99°95 7.692 
M983077 52°54.092' L010), 125) 
M983079 52°47.205' 100°58.041' 
M983073 SL°55,587 100°09.384' 
M993036 31236.570' 99°51.542' 
M993017 52°46.802' 100°57.852' 
M983074 52°00.408' 100°08.480' 
M993041 5202191" 100°09.754' 
M003007 52°S7.D07 LOMO 1299! 
M993032 ap hago fo i) 9 Ih 100°09.068' 
M993021 52°43.719' 100°40.582' 
M993033 51°50:039' 100°07.305' 
M993022 52°42.632' 100°41.958' 
M993039 51°48.000' 99°43.792' 
M003001 51°34.591' 99°41.305' 
M993024 52°44.569' 100°44.862' 
M993015 52°46.925° 100°53.561' 
M993012 32°93.878' 101°02.030' 


Vol. 119 

TDS* Cl SO, Be 

mg/I mg/1 mg/l ug/I pH 
64510 35700 3460 1700 6.87 
63488 35300 3220 1900 6.70 
61637 34300 3200 1200 6.75 
61327 33700 3170 1100 6.50 
61015 35400 1130 1700 6.29 
60300 33000 3170 1900 6.35 
59404 32800 3180 1100 6.70 
58646 32500 2970 2500 6.60 
58197 32100 2990 2300 6.64 
56545 31500 2780 2000 6.78 
55861 31300 2840 2700 6.73 
55855 31200 2770 2100 6.56 
55626 31000 2820 <150 6.34 
54825 31000 2800 2100 6.97 
54543 30200 2790 <150 7.40 
49068 27400 2600 <150 6.99 
48522 27200 2400 1900 6.86 
43680 22300 4590 <150 o:30 
42405 23700 2100 3200 708 
38566 21500 2090 1000 6.91 
37208 19900 3000 <150 8.38 
37189 18900 3690 240 7.99 
35856 18900 3010 <150 7.96 
32786 18500 1610 1000 6.98 
32356 18100 1620 110 7.88 
28857 15500 2300 2400 Tag 
28765 14400 3360 <150 15 
23793 12700 1080 340 6.48 
22319 11700 1400 2300 7.40 
21648 10800 2250 <150 6.94 
21100 12100 1011 <150 6.87 
20096 10300 1500 <150 5.98 
18236 9970 899 <150 762 
17453 8320 1580 <150 7.84 
16946 9730 808 <150 7.10 
14371 1400 2460 <150 189 
11914 5530 1790 390 6.93 

7189 3720 321 <150 8.28 

5892 2850 el <150 6.62 

5027 2490 180 <150 7.75 


“Hypersaline springs with 2 percursa 
> Hypersaline springs with algal mats, possibly 2 percursa 
© Total dissolved solids (TDS) 


having two (Celan 1979). Although the thalli are usu- 
ally biseriate, it is common to find some thalli that are 
uniseriate in part. Cells in the material collected from 
the saline springs bordering Lake Winnipegosis are a 
typical size (Abbott and Hollenberg 1976), between 
10 — 18 um wide and 24 — 28 um long. The thalli are 
light green because of the chloroplasts, but the cells 
from this site are often covered by amorphous rust- 
coloured clusters that can impart a brown, to rust, to 
orange, to red colour to the mats as a whole. This has 
led to the illusion that these mats are dominated by red 
and green algae (Bezys et al. 1997; Wadien 1984), when 
in fact only green algae (Chlorophyta) are present. 
This species is usually found with other algae, fre- 
quently forming floating entangled masses (often called 


green tide or sea lettuce) in upper tidepools and salt 


marshes. In North America, this taxon 1s found along © 


the Pacific coast (Abbott and Hollenberg 1976; Set- 
chell and Gardner 1920, Garbary et al. 1985), the At- 
lantic coast (South 1984; Wynne 1986), and in the Arc- 


tic (Setchell and Gardner 1920). Typically a marine | 


organism, there is one previous report that P percursa 
may also occur inland in environments where salt con- 
centrations are elevated (Kornmann 1956). Although 
reported in many locations, this filamentous alga usu- 
ally does not occur in large quantities, and is generally 


a minor contributor to blooms dominated by closely- | 
related species of Cladophora and Enteromorpha — 


(Hayden and Waaland 2002; Setchell and Gardner 


1920; Taylor et al. 2001). In the hypersaline springs _ 


2005 


LONDRY, BADIOU, AND GRASBY: IDENTIFICATION OF A MARINE GREEN ALGA 85 


ft * > hee 
tS 3 
*e +» aie 


CT Sy 
ee tse ‘ om 


* 


etre aoe 


FIGURE 2. Photograph of a hypersaline spring containing extensive mats of P. percursa. A 15 cm tent peg inserted beside the 


spring is included for scale. 


near Lake Winnipegosis, it was by far the dominant 
organism, associated with various pennate diatoms 
and cyanobacteria (including members of the Oscil- 
latoriaceae) (unpublished data) but not other filamen- 
tous green algae. 


Habitat and distribution 

There are numerous saline springs along the western 
shore of Lake Winnipegosis (Grasby 2000; Bezys et al. 
1997; McKillop et al. 1992) as well as in north-eastern 
Alberta (Grasby and Chen 2005), however, prolific 
growth of 2 percursa has only been confirmed at the 
Dawson Bay salt springs located on the north-western 
shore of the lake. Similar algal growths have been 
observed at other sites west of Pelican Bay and south 
of Red Deer River, but detailed analyses have not been 
done to confirm ? percursa. Salinity at these sites is 
typically greater than that of the springs located fur- 
ther south in Manitoba, and this may explain the dom- 
inance of 2 percursa. The salinity of the springs is 
thought to be increasing over time (Patterson et al. 
1997), and although the full range salinity tolerance is 
unknown, it does not seem to be limiting growth of this 
organism, but rather promoting it. 2 percursa can 
grow in diluted seawater over a range of 0-34 PSU 
(practical salinity units), with an optimum of 24 PSU 
(Taylor et al. 2001), but tolerance above seawater has 
not been tested. 2 percursa has a broad tolerance to 
environmental conditions of irradiance (9-175 pmol 
ms!) and temperature (at least 10-30°C), as well as 


FIGURE 3. Photomicrograph (1000x) of filaments of Percur- 
sara percursa from a hypersaline spring west of Lake 
Winnipegosis. The biseriate cells are paired in two 
longitudinal rows in each filament. Note the large ac- 
cumulation of material on the right strand that imparts 
a rust-red color to the mats as a whole, although the 
filaments themselves are green. The scale bar repre- 
sents 50 um. 


86 THE CANADIAN FIELD-NATURALIST 


N and P concentrations [1-200 uM PO,*, 10-1000 uM 
NG; 1-100 uM NH,"] (Taylor et al. 2001). Growth 
of “green tide” algae like P pervcursa is generally en- 
hanced by eutrophic conditions, but the springs west 
of Lake Winnipegosis are pristine discharges, with 
the only potential sources of significant amounts of 
N and P from sporadic incidences of bird defecation. 

The distribution of P percursa was compared to 
water chemistry parameters by principle component 
analysis (PCA) using The Unscrambler® 7.6, a multi- 
variate statistical software package. The first two axes 
of the PCA explained 86% of the total variance and 
revealed differences in the water chemistry between 
hypersaline springs with and without mat communi- 
ties (Figure 4). PCA axis | explained 65% of the total 
variation between the springs and clearly separated 
hypersaline springs with mat communities from those 
without. Total dissolved solids (TDS = the sum of 
major cations and anions) and chloride explained 95.2 
and 94.6% of the variance in the scores of the hyper- 
saline springs on axis 1, respectively. All hypersaline 
springs with mat communities were situated to the 
right of axis | and positively correlated to TDS and 
chloride, indicating that high salinities are required 
for the growth of P percursa. Tron and sulphate con- 
centrations also appeared to be correlated to the pres- 
ence of mats and explained 49.0 and 56.5% of the 


PC2 


-1.0 -0.5 


Vol. 119 


variance on axis 1, respectively. Additionally, PCA 
axis 2 explained 21% of the variation between the 
hypersaline springs. On this axis pH was the most 
important factor, explaining 55.3% of the variance on 
axis 2, and indicated that the growth of P percursa 
occurred in springs with lower pH. 

In addition to the above parameters, other factors 
must also determine the distribution, as not all springs 
at each site contain mats of P pervcursa. The temper- 
ature of the waters varied seasonally, ranging from 
5°C (temperature of the source water) up to 30°C in 
the summer, but densities did not correlate with tem- 
perature. Growth of the organism is restricted to springs 
with high flow rates that form open pools, and it is 
not observed in low-flowing seeps in the area even if 
the water chemistry is compatible. In addition, distinct 
pools or cauldrons just a few meters apart, or even 
touching, had extensive growth of mats in one pool 
and nothing in the other. Considering the motility of 
the species, and its widespread distribution in the area, 
lack of colonization would not explain the patchy 
distribution. There could be a subtle difference in the 
chemistry of the water at each outlet, although chemi- 
cal measurements to date indicate that all the water 
from these sites originates from the same source. There 
could be a difference in predation to explain the dis- 
parity of mat development in different springs. Studies 


1.0 


0.0 0.5 


PC1 


FIGURE 4. Principle component analysis (PCA) ordination biplot displaying grouping of hypersaline springs with respect to 
water chemistry. The symbols represent springs without 2 percursa (closed circles), with mat communities not yet 
confirmed as ? percursa (open circles), and with confirmed mats of 2 percursa (open stars). The inset table lists the rela- 
tive contributions (in %) of each chemical parameters to the variance on each of the two axes. 


2005 


of rotifer populations in areas containing P. percursa 
found that this predator 1s limited by high salt concen- 
trations as well as higher temperatures (Saunders- 
Davies 1998). The dynamics of growth and predation 
require further study at this site. 


Further research 

The ability of P percursa to grow and thrive in 
this extreme environment certainly merits further sci- 
entific investigation. The extent to which this domi- 
nant primary producer supports an entire ecosystem 
of microbes in the mat community is the subject of 
ongoing research. This organism supports a dynamic 
community of anaerobes including sulphate-reducing 
bacteria and methanogens. The potential for methano- 
genic fermentation of 2 percursa biomass at lower 
salinities has been demonstrated previously, and pro- 
posed as a future mechanism for the production of a 
renewable source of biogas (Schramm and Lehnberg 
1984). The extent to which this organism affects the 
precipitation and mineralization of carbonates (calci- 
um carbonate) and iron (hematitic- to limonitic) at 
these sites (Bezys et al. 1997), causing the distinctive 
reddish brown sinter or tufa mounds of springs on the 
salt pans, also requires further investigation. These 
mats also likely support a unique community of inver- 
tebrates and epiphytic algae, which to date have not 
been investigated to our knowledge. 


Acknowledgments 

This research was supported by NSERC. Thanks 
to K. Dawson, H. Grover, and S. Mohammed for col- 
lection of samples. Thanks to H. Kling at the Fresh- 
water Institute (Winnipeg, Manitoba) for helpful advice 
regarding the identification of P percursa. 


Literature Cited 

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Bezys, R. K., E. B. Ducharme, J. D. Bamburak, and M. 
A. F. Fedikow 1997. A geochemical study of saline brine 
sediments as a guide to Prairie-type microdisseminated 
mineralization and other precious metals in west-central 
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Bliding, C. 1963. A critical survey of European taxa in Ulvales 
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Burchill, C. A., and N. C. Kenkel. 1991. Vegetation-envi- 
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Celan, M. 1979. Deux algues vertes (Chlorophyceés) nou- 
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Garbary, D. J., G. I. Hansen, and R. F. Scagel. 1985. 
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Grasby, S. E. 2000. Saline spring geochemistry, west-cen- 
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Manitoba Industry, Trade and Mines, Manitoba Geologi- 
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ton basin in response to Pleistocene glaciation: Geology 
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Grasby, S. E. and Z. Chen. 2005. Subglacial recharge into 
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Hayden, H. S., and J. R. Waaland. 2002. Phylogenetic 
systematics of the Ulvaceae (Ulvales, Ulvophyceae) using 
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cology 38: 1200-1212. 

Kornmann, P. 1956. Zur Morphologie und Entwicklung 
von Fercursaria percursa. Helgolander Wissenschaftliche 
Meeresuntersuchungen 5: 259-272. 

McKillop, W. B., R. T. Patterson, L. D. Delorme, and T. 
Nogrady. 1992. The origin, physico-chemistry and biotics 
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O’Kelly, C. J., B. Wysor, and W. K. Bellows. 2004. Gene 
sequence diversity and the phylogenetic position of algae 
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Patterson, R. T., W. B. McKillop, S. Kroker, E. Nielsen, and 
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Received 5 July 2004 
Accepted 25 February 2005 


New Distribution Records and Biogeography of Ca//igrapha Species 
(Leaf Beetles), in North America (Coleoptera: Chrysomelidae, 


Chrysomelinae) 


JESUS GOMEZ-ZURITA 


Area de Biologia Animal, Departamento de Zoologia y Antropologfa Fisica, Universidad de Murcia, 30071 Murcia, Spain. 


Gomez-Zurita, Jesis. 2005. New distribution records and biogeography of Ca//igrapha species, (Leaf Beetles), in North 
America (Coleoptera: Chrysomelidae, Chrysomelinae). Canadian Field-Naturalist 119(1): 88-100. 


The study of a large number of collection specimens from the US National Museum of Natural History belonging to the 
genus Ca/ligrapha Chevrolat (Chrysomelidae) from the Nearctic region has provided with one new province record and one 
new species record for Canada and 63 new US State records for a total of 20 taxa. These new records usually correspond to 
adjacent areas to already identified ranges of distribution, expanding considerably the known geographical boundaries of 
the species studied. The zoogeography of the genus Ca//igrapha in North America is discussed based on the updated infor- 


mation on the distribution of the genus. 


Key Words: Ca//igrapha, Leaf beetles, Nearctic region, zoogeography, biodiversity, distribution, new records. 


The New World chrysomeline genus Ca//igrapha 
Chevrolat 1837 (Chrysomelidae) is currently recog- 
nized to have more than 80 species and subspecies 
distributed from Alaska and all Canada provinces (and 
Northwest Territories) to Argentina in South America 
(Blackwelder 1982; Arnett et al. 2002). Approximately 
one third of these species are present in North America 
north of Mexico (Arnett 1968; Arnett et al. 2002), the 
biogeographical region for which we have a better 
knowledge of the taxonomy and also geographical 
ranges of the species. This group is best known taxo- 
nomically and biologically from Eastern Canada, 
thanks to the observations of W. J. Brown (1940, 1945, 
1958) and J. G. Robertson (1966), who studied in detail 
their distribution along with trophic preferences and 
their cytology. But even for a region like North Amer- 
ica and a well-studied zoological group like the Chry- 
somelidae, information about distribution is usually 
scattered in catalogues of regional fauna (e.g., Blatch- 
ley 1910; Balsbaugh and Hays 1972; Clark 2000) or 
in the form of far from complete State records in more 
general species accounts (e.g., Wilcox 1975; Downie 
and Arnett 1996; Riley et al. 2003). Besides, our under- 
standing about species distributions can be in partic- 
ular cases quite confuse due to unreliable taxonomic 
designations for morphologically very similar species. 
Problematic species identification in Ca//igrapha is 
rather common for several closely related taxa be- 
cause it requires information about host plants, most of 
the times lacking from collection specimens, or large 
series of individuals to capture the species variation 
within the species. 

In order to characterize in higher detail the geo- 
graphical ranges of the species of Ca//igrapha in 
North America it is still required extensive field work 
in new and already prospected areas, paying particu- 


lar attention to the plants to which the beetles appear 
associated in the field. Furthermore, the study of the 
collections in Museums has logically enormous poten- 
tial in providing this type of information, with new 
and interesting data to be gathered. Both sources of 
information are used in this work, together with pub- 
lished data, to present an updated account of the infor- 
mation about distribution for 20 taxa of Ca//igrapha 
in North America for which several new state records 
have been found. In most of the cases, these new 
records correspond to regions adjacent to the already 
identified areas of distribution of each taxon, broad- 
ening the known distributional ranges of these species. 
The study of a single, but large, source of Museum 
material like that deposited in the Smithsonian Insti- 
tution has proved very fruitful in providing with new 
data. Nevertheless, it is possible that the investigation 
of the accessions and undetermined specimens from 
other Museums, particularly in Canadian and US Insti- 
tutions, will certainly add new useful information 
about the distribution of these beetles. 


Materials and Methods 
Specimens studied 

Collection data for some specimens were obtained 
by the author in two entomological campaigns in north- 
eastern North America in the summer of 2001 and 
eastern Canada in 2002. Field work done independ- 
ently by Daniel J. Funk in the same areas also provid- 
ed with important new distribution records. A large 
number of accessions from the collection at the US 
National Museum of Natural History (NMNH; Smith- 
sonian Institution, Washington, DC) were also studied, 
which included 2030 specimens belonging to the genus 
Calligrapha. 


88 


2005 


Species identification 

All the specimens reported here and all the other 
museum accessions available were personally identi- 
fied by the author of this work, who has collected and 
investigated the taxonomy and phylogeny of the genus 
Calligrapha since 2000. The museum accessions 
were sorted out separating all the species that can be 
easily identified at a glance because of their charac- 
teristic features, such as the species belonging to the 
subgenera 4idensomela, Coreopsomela and Graphi- 
callo, or the Calligrapha s. str species C. apicalis, C. 
ignota, C. rowena, and others. The other samples and 
every single specimen reported in this work were key 
out using the comprehensive keys for North Ameri- 
can Calligrapha in Brown (1945) and Wilcox (1972), 
but were also compared with correctly identified mate- 
rial in the British Natural History Museum Entomology 
collection and in the reference specimen and picture 
collection of the author. Moreover, the individual speci- 
mens were always compared with the original descrip- 
tion of the species to further confirm the identity. The 
over 2000 identified specimens were labeled with their 
taxonomic assignment on white cardboard labels in- 
cluding “J. G6mez-Zurita det.” and the year when the 
identification was done. The comparatively few spec- 
imens collected by D. J. Funk and other colleagues 
for the author’s research on the molecular phylogeny 
of the genus are temporally held in the author’s col- 
lection before the studies are finished and their final 
depository decided. The remaining specimens belong 
and are deposited in the US NMNH collection. 


Data presentation 

For each Ca//igrapha species, distribution informa- 
tion is given by state for published records. Full details, 
including county and specific locality data when known, 
collection date, the name of the collector, as well as 
the number of specimens studied, are given in those 
cases where a new state record has been registered. 
Similar distributional information is presented graph- 
ically on a series of maps where references from the 
literature and new records are distinguished (Figure | 


a-p). 


Results 


Calliigrapha alni Schaeffer 1928 (Figure 1a) 

Canada: Alberta (Brown 1945; Robertson 1966; Bousquet 
1991; Riley et al. 2003), Manitoba (Brown 1945; Robert- 
son 1966; Bousquet 1991; Riley et al. 2003), New Bruns- 
wick (Bousquet 1991; Riley et al. 2003), Nova Scotia 
(Schaeffer 1928; Brown 1945; Robertson 1966; Wilcox 
1972, 1975; Bousquet 1991; Cavey 1994; Downie and 
Arnett 1996; Riley et al. 2003), Ontario (Schaeffer 1928; 
Robertson 1966; Bousquet 1991; Cavey 1994; Riley et al. 
2003), Quebec (Robertson 1966; Bousquet 1991; Downie 
and Arnett 1996; Riley et al. 2003), Saskatchewan (Rob- 
ertson 1966; Bousquet 1991; Riley et al. 2003). U.S.A.: 
Maine (Schaeffer 1928; Cavey 1994; Downie and Arnett 
1996; Riley et al. 2003), Maryland (Cavey, 1994; Downie 
and Arnett 1996; Riley et al. 2003), Massachusetts (Schaef- 
fer 1928; Cavey 1994; Riley et al. 2003), Michigan (Wilcox 


GOMEZ-ZURITA: NEW RECORDS AND BIOGEOGRAPHY OF C4ZLL/GRAPHA 89 


1972, 1975; Cavey 1994; Downie and Arnett 1996; Riley 
et al. 2003), Minnesota (Riley et al. 2003), New Jersey 
(Schaeffer 1933; Cavey 1994; Riley et al. 2003), New York 
(Schaeffer 1933; Cavey 1994; Downie and Arnett 1996; 
Riley et al. 2003), Ohio (Wilcox 1954; Cavey 1994; Riley 
et al. 2003), Vermont (Schaeffer 1928; Cavey 1994; Riley 
et al. 2003), West Virginia (Wilcox 1972, 1975; Cavey 
1994; Downie and Arnett 1996; Clark 2000; Riley et al. 
2003), Wisconsin (Cavey 1994; Riley et al. 2003). 

New HAMPSHIRE: Coos Co., Jefferson, 4 May 1936, 
A. E. Brower collector (1) 


Calligrapha alnicola Brown 1945 (Figure |b) 

Canada: New Brunswick (Brown 1945; Robertson 1966; 
Bousquet 1991; Clark 2000; Riley et al. 2003), Nova Sco- 
tia (Bousquet 1991; Riley et al. 2003), Ontario (Brown 
1945; Robertson 1966; Wilcox 1972, 1975; Bousquet 1991; 
Downie and Arnett 1996; Riley et al. 2003), Quebec 
(Brown 1945; Robertson 1966; Wilcox 1972, 1975; Bous- 
quet 1991; Downie and Arnett 1996; Riley et al. 2003). 
U.S.A.: Kentucky (Clark 2000; Riley et al. 2003), Maine 
(Clark 2000; Riley et al. 2003), Michigan (Clark 2000; 
Riley et al. 2003), New York (Riley et al. 2003). 

MINNESOTA: Cook Co., route 61, North, 47°56.879'N 
89°41.875'W, 14 August 2004, on A/zus sp., D. J. 
Funk collector (2) 

New HAmpPSHIRE: Grafton Co., Hanover, K. W. Cooper 
coll. (1); Strafford Co., Durham, 12 June 1907 (1) 
VERMONT: Bennington Co., East Dorset, 8 June 1959, 

C. T. Parsons coll. (1); Orleans Co., near Barton, 
16 July 2001, on A/zus sp., D. J. Funk collector (1) 
WEST VIRGINIA: Tucker Co., Canaan Valley Resort 
S. P., 18 July 2001, J. Gomez-Zurita and D. P. Duran 
collector (1) 
WISCONSIN: Bayfield Co., Bayfield, Wickham col- 
lector (1) 


Caligrapha amator Brown 1945 (Figure 1c) 

Canada: Ontario (Brown 1945; Robertson 1966; Wilcox 1972, 
1975; Bousquet 1991; Downie and Arnett 1996; Riley et 
al. 2003). U.S.A.: Wyoming (Lawson 1976; dubious ac- 
cording to Riley et al. 2003). 

QUEBEC: Quebec, Maisonneuve, 25 August 1901, C. 
Stevenson collector (3); St. Lawrence River, St. 
Regis, on 7i/ia americana, C. O. Houghton collec- 
tor (1) 


Caliigrapha amelia Knab 1909 (Figure |c) 

U.S.A.: Alabama (Balsbaugh and Hays 1972; Riley et al. 
2003), District of Columbia (Knab 1909; Leng 1920; Riley 
et al. 2003), Maryland (Knab 1909; Riley et al. 2003), 
New Jersey (Knab 1909; Wilcox 1972, 1975; Riley et al. 
2003), New York (Knab 1909; Leng 1920; Leonard 1926; 
Wilcox 1972; Downie and Arnett 1996; Riley et al. 2003), 
Ohio (Wilcox 1954, 1972, 1975: Downie and Arnett 1996; 
Riley et al. 2003), Rhode Island (Sikes 1999*; Riley et 
al. 2003), South Carolina (Kirk 1970; Riley et al. 2003), 
Virginia (Knab 1909; Leng 1920; Wilcox 1972, 1975; 
Downie and Arnett 1996; Riley et al. 2003), West Virginia 
(Knab 1909; Clark 2000; Riley et al. 2003). 

CONNECTICUT: New London Co., Lyme, 28 May 
1918, W. S. Fisher collector, on A/zus sp. (1) 


90 THE CANADIAN FIELD-NATURALIST Vol. 119 


FIGURE |. Geographical distributions of several Ca//igrapha species in North America. The shaded areas represent published 
province (Canada) or US State records and the symbols correspond to the new province or state records reported in this 
work for C: a/ni(a), C. alnicola(b), C. amator and C. amelia(c), C. apicalis (a), C. bidenticola(e), C. californica s. 1. (f), 
C. confluens (g), C. ignota (h). 


DELAWARE: Sussex Co., Rehoboth, 6 April 1943, G. Caligrapha apicalis Notman 1919 (Figure 1d) 
H. Dieke collector, in beach washup (1) Canada: Ontario (Brown 1945; Robertson 1966; Bousquet 
GeorGIA: De Kalb Co., 5 May 1951, H. R. Dodge 1991; Riley et al. 2003), Quebec (Brown 1945; Robert- 


collector (1); Spaulding Co., E . t, 29 March son 1966; Wilcox 1972, 1975; Bousquet 1991; Downie 
183 7e0n in pote ce Coen hy and Arnett 1996; Riley et al. 2003). U.S.A.: Michigan 


E (Wilcox 1972, 1975; Downie and Arnett 1996; Riley et 
MASSACHUSETTS: F. Knab collector (1) al. 2003), New York (Notman 1919; Leonard 1926; 
PENNSYLVANIA: Cumberland Co., Mount Holly Wilcox 1972, 1975; Downie and Arnett 1996; Riley et al. 
Springs, | September 1918, R. M. Fonts collector 2003). 
(1); Dauphin Co., Heckton Mills, 8 July 1910, P. MaryLanp: Calvert Co., Plum Point, 28 May 1922, 
R. Myers collector (1) L. L. Buchanan collector (1) 


2005 


x SAUER nant 
* Calligraph bloat) 


GOMEZ-ZURITA: NEW RECORDS AND BIOGEOGRAPHY OF C4ZL/GRAPHA 9] 


FiGureE |. (Continued). C. kzabiand C. rhoda (i), C. ostryae (j), C. philadelphica(k), C. rowena (\), C. scalaris (m), C. spiraea 
(n), C! suturella and C. sylvia (0), and C. verrucosa and C. vicina (p). The symbols are approximately placed on the locali- 
ties in the collection data. Larger symbols are used to identify US State records without specific locality. 


Caliigrapha bidenticola Brown 1945 (Figure le) 

Canada: New Brunswick (Brown 1945; Robertson 1966; 
Wilcox 1972, 1975; Downie and Arnett 1996; Riley et al. 
2003), Ontario (Brown 1945; Robertson 1966; Bousquet 
1991; Riley et al. 2003), Quebec (Brown 1945; Robertson 
1966; Bousquet 1991; Riley et al. 2003). U.S.A.: Alabama 
(Balsbaugh and Hays 1972; Wilcox 1972; Riley et al. 
2003), Arkansas (Riley et al. 2003), Colorado (Leng 1920; 
Wilcox 1972, 1975; Downie and Arnett 1996; Riley et al. 
2003), Connecticut (Britton 1920; Brown 1945; Riley et 
al. 2003), District of Columbia (Riley et al. 2003), Florida 


(Monroés 1955; Wilcox 1972, 1975; Riley et al. 2003), 
Georgia (Riley et al. 2003), Illinois (Riley et al. 2003), 
Indiana (Blatchley 1910; Leng 1920; Downie and Arnett 
1996; Riley et al. 2003), Iowa (Riley et al. 2003), Kansas 
(Douglass 1929; Riley et al. 2003), Kentucky (Riley et al. 
2003), Louisiana (Riley et al. 2003), Maine (Riley et al. 
2003), Maryland (Riley et al. 2003), Massachusetts (Downie 
and Arnett 1996; Riley et al. 2003), Missouri (Riley and 
Enns 1979; Riley et al. 2003), Nebraska (Powell 1932; 
Riley et al. 2003), New Jersey (Riley et al. 2003), New York 
(Leonard 1926; Downie and Arnett 1996; Riley et al. 2003), 


92 THE CANADIAN FIELD-NATURALIST 


Ohio (Hughes 1944; Wilcox 1954; Riley et al. 2003), Okla- 
homa (Shaddy and Drew 1967), Pennsylvania (Riley et al. 
2003), Rhode Island (Davis 1904; Sikes 1999*; Riley et al. 
2003), South Dakota (Johnson 1930; Kirk and Balsbaugh 
1975; Riley et al. 2003), Tennessee (Riley et al. 2003), 
Texas (Wilcox 1972, 1975; Downie and Arnett 1996; Riley 
et al. 2003), West Virginia (Clark 2000; Riley et al. 2003), 
Wisconsin (Riley et al. 2003). 

DELAWARE: Dewey Beach to Maryland, Assateague 
Park, 1 July 1985, dead on beach, P. Schaefer collec- 
tor (1) 

MICHIGAN: E. S. George Res., Pinckney, 8 June 1947, 
B. Summerville collector (2) - 


Caliigrapha cakifornica s. £ (Linell 1896) (Figure 1f) 

Canada: Alberta (Beller and Hatch 1932; Brown 1945; Bous- 
quet 1991; Riley et al. 2003), British Columbia (Brown 
1945; Hatch 1971; Wilcox 1972, 1975; Bousquet 1991; 
Riley et al. 2003), Manitoba (Brown 1945; Bousquet 1991; 
Riley et al. 2003), New Brunswick (Brown 1945; Bousquet 
1991; Riley et al. 2003), Northwest Territories (Bousquet 
1991; Riley et al. 2003), Nova Scotia (Beller and Hatch 
1932; Brown 1945; Riley et al. 2003), Ontario (Morris 
1914; Bousquet 1991; Riley et al. 2003), Prince Edward 
Island (Riley et al. 2003), Quebec (Brown 1945; Robert- 
son 1966; Bousquet 1991; Riley et al. 2003), Saskatchewan 
(Brown 1945; Bousquet 1991; Riley et al. 2003). U.S.A.: 
California (Leng 1920; Beller and Hatch 1932; Schaeffer 
1933; Brown 1945; Monrés 1955; Wilcox 1972, 1975; 
Riley et al. 2003), Colorado (Beller and Hatch 1932; Riley 
et al. 2003), Connecticut (Britton 1920; Brown 1945; Riley 
et al. 2003), District of Columbia (Beller and Hatch 1932: 
Riley et al. 2003), Idaho (Schaeffer 1933; Hatch 1971), 
Indiana (Blatchley 1910; Leng 1920; Beller and Hatch 
1932; Riley et al. 2003), Iowa (Riley et al. 2003), Kansas 
(Douglass 1929; Riley et al. 2003), Maine (Wilcox 1972, 
1975; Downie and Arnett 1996; Riley et al. 2003), Mary- 
land (Riley et al. 2003), Massachusetts (Riley et al. 2003), 
Michigan (Lawson 1976; Riley et al. 2003), Minnesota 
(Riley et al. 2003), Montana (Hatch 1971; Riley et al. 
2003), Nebraska (Powell 1932; Riley et al. 2003), New 
Jersey (Riley et al. 2003), New York (Leonard 1926; Law- 
son 1976; Riley et al. 2003), North Dakota (Wilcox 1972, 
1975; Downie and Arnett 1996), Ohio (Hughes, 1944; 
Wilcox 1954; Riley et al. 2003), Oregon (Hatch 1971), 
Pennsylvania (Riley et al. 2003), Rhode Island (Davis 
1904; Sikes 1999*; Riley et al. 2003), South Carolina (Riley 
et al. 2003), South Dakota (Johnson 1930; Wilcox 1972, 
1975; Kirk and Balsbaugh 1975; Riley et al. 2003), Utah 
(Riley et al. 2003), Virginia (Williams 1989; Riley et al. 
2003), Washington (Beller and Hatch 1932; Hatch 1971; 
Riley et al. 2003), West Virginia (Clark 2000; Riley et al. 
2003), Wisconsin (Riley et al. 2003), Wyoming (Lawson 
1976; Riley et al. 2003). 

ILLINOIS (C. c. coreopsivora): Cook Co., Chicago, 30 
May 1902, on the beach, A. B. Wolcott collector (1); 
Cook Co., Chicago, | June 1904, A. B. Wolcott coll. 
(1) 

VERMONT (C. c. coreopsivora): Bennington Co., East 
Dorset, 8 July 1959, C. Parsons collector (12); 
Chittenden Co., Colchester, Colchester Pond, 9 May 
1975, M. Langworthy collector (1); LaPlatt River, 
20 September 1979, B. J. Norman collector (1) 


Vol. 119 


Caliigrapha confiuens Schaeffer 1928 (Figure 1g) 

Canada: New Brunswick (Bousquet 1991; Riley et al. 2003), 
Nova Scotia (Schaeffer 1928: Brown 1945; Robertson 
1966; Wilcox 1972, 1975; Bousquet 1991; Downie and 
Arnett 1996; Riley et al. 2003), Ontario (Brown 1945; 
Robertson 1966; Bousquet 1991; Riley et al. 2003), Que- 
bec (Brown 1945; Robertson 1966; Bousquet 1991; Riley 
et al. 2003). U.S.A.: Maine (Schaeffer 1928; Riley et al. 
2003), Massachusetts (Schaeffer 1928; Wilcox 1972, 1975; 
Downie and Arnett 1996; Riley et al. 2003), Nebraska 
(Powell 1932; reported as dubious by Riley et al. 2003), 
New York (Brown 1945; Riley et al. 2003), Virginia (Brown 
1945), West Virginia (Clark 2000; Riley et al. 2003). 

MINNESOTA: Cass Co., Gull Lake, 3 June 1925, F 
Uhler collector (1) 

NEw HAmpsSHIRE: Mt. Plsnt. Hse., July, A. Feynes 
collection (1) 

PENNSYLVANIA: Erie Co., North East, 4 July 1916, on 
Alnus, R. A. Cushman collector (1) 

VERMONT: Chittenden Co., Westford, 1 October 1968, 
D. L. Pouliot collector (1); Caledonia Co., nr Shef- 
field, 20 August 1999, on A/nzus sp., Daniel J. Funk 
collector (1) 


Caliigrapha ignota Brown 1945 (Figure 1h) 

Canada: Manitoba (Bousquet 1991; Riley et al. 2003), New 
Brunswick (Brown 1945; Bousquet 1991; Riley et al. 
2003), Nova Scotia (Brown 1945; Wilcox 1972, 1975; 
Bousquet 1991; Downie and Arnett 1996; Riley et al. 
2003), Ontario (Brown 1945; Bousquet 1991; Riley et al. 
2003), Prince Edward Island (Brown 1945; Riley et al. 
2003), Quebec (Riley et al. 2003). U.S.A.: Maine (Riley 
et al. 2003), Minnesota (Wilcox 1972, 1975; Downie and 
Arnett 1996; Riley et al. 2003), North Carolina (Riley et 
al. 2003), Ohio (Riley et al. 2003), Pennsylvania (Linnell 
1896; Brown 1945; Wilcox 1972; Downie and Arnett 1996; 
Riley et al. 2003), Tennessee (Wilcox 1972, 1975; Downie 
and Arnett 1996; Riley et al. 2003), West Virginia (Clark 
2000; Riley et al. 2003). 

MICHIGAN: Allegan Co., Macatawa B’h, 10 June 1906, 
A. B. Wolcott collector (2); Marquette Co., Mar- 
quette, June 1928, Van Dyke Collection (2) 

NEw York: Erie Co., Buffalo, E. P. V. collector, Col- 
lection Ashmead (1); Long Island, 18 April 1903, 
Sherman Collection (1); Long Island, Sherman Col- 
lection (4); New York, Sherman Collection (2) 


Caliigrapha knabi Brown, \940 (Figure 1i) | 

Canada: Manitoba (Bousquet 1991; Riley et al. 2003), Ontario 
(Bousquet 1991; Riley et al. 2003), Quebec (Brown 1940, 
1945; Wilcox 1972, 1975; Bousquet 1991; Downie and 
Arnett 1996; Riley et al. 2003). 

MARYLAND: Prince George’s Co., Accokeek, 24 April | 
2003, on Cornus amomum, C. L. & S. L. Staines | 
collector (2) 

NorTH CAROLINA: Jackson Co., Balsam, 25 June 
1973, Rosenberg Collection (1) 


Caliigrapha ostryae Brown 1945 (Figure 1j) 

Canada: Ontario (Brown 1945; Robertson 1966; Wilcox 1972, 
1975; Bousquet 1991; Downie and Arnett 1996; Riley et al. 
2003), Quebec (Brown 1945; Robertson 1966; Riley et al. 


2005 


2003). U.S.A.: Michigan (Brown 1945; Robertson 1966; 
Wilcox 1972, 1975; Downie and Arnett 1996; Riley et al. 
2003), New York (Brown 1945; Robertson 1966; Wilcox 
1972, 1975; Downie and Arnett 1996; Riley et al. 2003). 

PENNSYLVANIA: Pennsylvania, F. Knab Collection (1) 

WISCONSIN: Milwaukee Co., Milwaukee, Milwaukee 
Public Museum, F. Knab Collection (1); Milwaukee 
Co., 20 July 1900, F. R. collector, F. Knab Collec- 
tion (2) 


Caliigrapha philadelphica (Linnaeus 1758) (Figure 
Ik) 

Canada: British Columbia (Brown 1945; Robertson 1966; 
Hatch 1971; Wilcox 1972, 1975; Lawson 1976; Bousquet 
1991; Downie and Arnett 1996; Riley et al. 2003), Mani- 
toba (Brown 1945; Robertson 1966; Bousquet 1991; 
Downie and Arnett 1996; Riley et al. 2003), New Bruns- 
wick (Brown 1945; Robertson 1966; Wilcox 1972, 1975; 
Bousquet 1991; Downie and Arnett 1996; Riley et al. 2003), 
Nova Scotia (Bousquet 1991; Riley et al. 2003), Ontario 
(Morris 1914; Brown 1945; Robertson 1966; Bousquet 
1991; Riley et al. 2003), Quebec (Brown 1945; Robertson 
1966; Bousquet 1991; Downie and Arnett 1996; Riley et al. 
2003), Saskatchewan (Bousquet 1991; Riley et al. 2003). 
U.S.A.: Alabama (Balsbaugh and Hays 1972; Riley et al. 
2003), Connecticut (Britton 1920; Riley et al. 2003), Dis- 
trict of Columbia (Riley et al. 2003), Georgia (Wilcox 
1972, 1975; Downie and Arnett 1996; Riley et al. 2003), 
Idaho (Hatch 1971; Downie and Arnett 1996), Indiana 
(Blatchley 1910; Leng 1920), Kentucky (Riley et al. 2003), 
Maine (Downie and Arnett 1996; Riley et al. 2003), 
Maryland (Schroder et al. 1996; Riley et al. 2003), Mass- 
achusetts (Riley et al. 2003), Michigan (Riley et al. 2003), 
Mississippi (Dozier 1921; Riley et al. 2003), Missouri 
(Rogers 1856; Riley et al. 2003), Montana (Hatch 1971; 
Riley et al. 2003), Nebraska (Linell 1896; Leng 1920; Pow- 
ell 1932; Wilcox 1972, 1975; Downie and Arnett 1996; 
Riley et al. 2003), New Jersey (Riley et al. 2003), New 
York (Leonard 1926; Downie and Arnett 1996; Riley et al. 
2003), North Carolina (Brimley 1938; Riley et al. 2003), 
Ohio (Hughes, 1944; Wilcox 1954; Downie and Arnett 
1996; Riley et al. 2003), Pennsylvania (Brown 1945; Riley 
et al. 2003), Rhode Island (Davis 1904; Sikes 1999*: 
Riley et al. 2003), South Carolina (Kirk 1970; Riley et al. 
2003), Virginia (Riley et al. 2003), Washington (Hatch 
1971; Downie and Arnett 1996), West Virginia (Clark 

| 2000; Riley et al. 2003). 

MINNESOTA: Kanabec Co., Mora, 20 June 1907, 27 
June 1907, 25 July 1907, 1 August 1907, R. A. Vick- 
ery collector (1, 2, 2;°1) 

New HAmpsuire: Grafton Co., Woodsville, 31 July 
2001, on Cornus stolonifera, J. Gomez-Zurita col- 

| lector (1); Grafton Co., Hanover, K. W. Cooper (6) 

Nort Dakota: Cass Co., Fargo, 20 July 1918, I. N. 

Gabrielson collector (1) 

VERMONT: Bennington Co., East Dorset, 28 June 1957, 

| C.T. Parsons collector (1) 


Calligrapha rhoda Knab 1909 (Figure 11) 

Canada: Manitoba (Bousquet 1991; Riley et al. 2003), Ontario 
| (Brown 1945; Bousquet 1991; Riley et al. 2003). U.S.A.: 
Connecticut (Britton 1920; Riley et al. 2003), Illinois (Knab 


GOMEZ-ZURITA: NEW RECORDS AND BIOGEOGRAPHY OF CAZL/GRAPHA 93 


1909; Riley et al. 2003), Indiana (Knab 1909; Blatchley 
1910; Leng 1920; Wilcox 1972; Riley et al. 2003), Kansas 
(Knab 1909; Leng 1920; Wilcox 1972, 1975; Downie 
and Arnett 1996; Riley et al. 2003), Maryland (Knab 1909; 
Wilcox 1972, 1975; Downie and Arnett 1996; Riley et al. 
2003), Massachusetts (Knab 1909; Downie and Arnett 
1996; Riley et al. 2003), Michigan (Knab 1909; Riley et al. 
2003), Minnesota (Knab 1909; Riley et al. 2003), Mis- 
souri (Knab 1909; Wilcox 1972; Riley et al. 2003), New 
Hampshire (Knab 1909; Leng 1920; Wilcox 1972, 1975; 
Downie and Arnett 1996; Riley et al. 2003), New Jersey 
(Riley et al. 2003), New York (Leonard 1926; Riley et al. 
2003), Rhode Island (Sikes 1999*; Riley et al. 2003), West 
Virginia (Clark 2000; Riley et al. 2003), Wisconsin (Knab 
1909; Leng 1920; Wilcox 1972, 1975; Downie and Arnett 
1996; Riley et al. 2003). 

Iowa: Johnson Co., Iowa City, May 1923, Wickham 
collector (1); Muscatine Co., Muscatine, 11 June 
1917, C. E. Smith collector (1); Iowa, Van Dyke 
Collection (1) 

LOUISIANA: Louisiana, F. Knab Collection (1) 

NEBRASKA: Cuming Co., West Point, June 1888, Koe- 
bele Collection (1) 

OKLAHOMA: Oklahoma Co., Harrah, 23 May 1916, 
W. D. Pierce collector (1); Payne Co., 1961, John 
F. Reinert collector (3) 


Calligrapha rowena Knab 1909 (Figure 11) 

Canada: Manitoba (Bousquet 1991; Riley et al. 2003), New 
Brunswick (Brown 1945; Robertson 1966; Bousquet 1991; 
Riley et al. 2003), Nova Scotia (Brown 1945; Robertson 
1966; Riley et al. 2003), Ontario (Knab 1909; Brown 1945; 
Robertson 1966; Bousquet 1991; Riley et al. 2003), Que- 
bec (Knab 1909; Brown 1945; Robertson 1966; Wilcox 
1972, 1975; Bousquet 1991; Downie and Arnett 1996; 
Riley et al. 2003). U.S.A.: Connecticut (Knab 1909; Brit- 
ton 1920; Riley et al. 2003), Georgia (Wilcox 1972, 1975; 
Downie and Arnett 1996; Riley et al. 2003), Indiana 
(Blatchley 1910; Leng 1920), Maine (Riley et al. 2003), 
Massachusetts (Knab 1909; Riley et al. 2003), Michigan 
(Wilcox 1972, 1975; Downie and Arnett 1996; Riley et al. 
2003), New Jersey (Riley et al. 2003), New York (Leonard 
1926; Brown 1945; Robertson 1966; Downie and Arnett 
1996; Riley et al. 2003), North Carolina (Brimley 1938; 
Riley et al. 2003), Ohio (Wilcox 1954; Riley et al. 2003), 
Pennsylvania (Knab 1909; Clark 2000; Riley et al. 2003). 

Iowa: Johnson Co., Iowa City, 4 July 1918, on Commus 
sp., L. W. Lindsey collector (1) 

KENTUCKY: Rockcastle Co., Pine Hill Cave, 27 March 
1967, Reddell and Andrews collector (1) 

MARYLAND: Garret Co., Oakland, 31 May 1942, 2 
June 1942, on Cornus sp., G. H. Dieke collector (1, 
1); Garret Co., Oakland, 6 June 1942, G. H. Dieke 
collector (1) 

MINNESOTA: Cedar Creek Natural History Area, 
45°40.000'N 93°15.000'W, 16 September 2004, on 
Cornus sp., D. J. Funk collector (1) 


Caliigrapha scalaris (e Conte 1824) (Figure 1m) 

Canada: Manitoba (Bousquet 1991; Riley et al. 2003), Ontario 
(Morris 1914; Brown 1945; Robertson 1966; Bousquet 
1991; Riley et al. 2003), Quebec (Brown 1945; Robertson 


94 THE CANADIAN FIELD-NATURALIST 


1966; Wilcox 1972, 1975; Bousquet 1991; Downie and 
Arnett 1996; Riley et al. 2003). U.S.A.: Alabama (Bals- 
baugh and Hays 1972; Riley et al. 2003), Arkansas (Riley 
et al. 2003), Connecticut (Britton 1920; Riley et al. 2003), 
District of Columbia (Riley et al. 2003), Florida (Le Conte 
1824; Leng 1920; Blatchley 1924; reported as dubious by 
Riley et al. 2003), Georgia (Wilcox 1972, 1975; Downie 
and Arnett 1996; Riley et al. 2003), Indiana (Blatchley 
1910; Leng 1920; Powell 1932; Downie and Arnett 1996), 
Kansas (Douglass 1929; Robertson 1966; Riley et al. 
2003), Maine (Riley et al. 2003), Massachusetts (Riley et 
al. 2003), Michigan (Downie and Arnett 1996), Missouri 
(Jacoby 1880-1888; Riley and Enns 1979), Nebraska 
(Rogers 1856; Linell 1896; Leng 1920; Powell 1932; 
Riley et al. 2003), New Jersey (Riley et al. 2003), New 
York (Le Conte 1824; Leonard 1926; Downie and Arnett 
1996; Riley et al. 2003), North Carolina (Brimley 1938; 
Riley et al. 2003), Ohio (Hughes 1944; Wilcox 1954; Riley 
et al. 2003), Oklahoma (Fenton 1944; Shaddy and Drew 
1967; Riley et al. 2003), Pennsylvania (Riley et al. 2003), 
Rhode Island (Davis 1904; Sikes 1999; Riley et al. 2003), 
South Carolina (Kirk 1969, 1970; Riley et al. 2003), South 
Dakota (Johnson 1930; Kirk and Balsbaugh 1975; Riley 
et al. 2003), Texas (Linell 1896; Leng 1920; Wilcox 1972, 
1975; Downie and Arnett 1996; Riley et al. 2003), West 
Virginia (Clark 2000; Riley et al. 2003). Mexico (Jacoby 
1880-1888; Blackwelder 1982). 

ILLINOIS: N. Illinois, F. Knab Collection (1) 

LOUISIANA: Franklin Co., Wisner, February 22-23, 
under bark of elm, R. A. St. George collector (2); 
Orleans Co., New Orleans, 20 January 1944, E. S. 
Ross collector (4) 

MINNESOTA: Clay Co., Moorhead, 3 July 1980, on 
American elm (1) 

Mississippi: Adams Co., Natchez, 15 June 1909, 17 
June 1909, 19 June 1909, 21 June 1909, in Spanish 
moss, E. S. Tucker collector (2, 1, 1, 5) 

NEw HAMPSHIRE: Carroll Co., Chocorua, altitude 
3470 ft., G. H. Dieke Collection (1); Hillsborough 
Co., Manchester, 18 May, 22 May, 9 June, 15 June, 
W. S. Abbott 1932 thru Bridwell (1, 1, 1, 1) 

WISCONSIN: Milwaukee Co., 30 July 1900, F. Rauter- 
berg collector (1); Milwaukee Co., F. Knab Collec- 
tion (1) 


Calliigrapha spiraea (Say 1826) (Figure |n) 

Canada: Ontario (Wheeler and Hoebeke 1979; Bousquet 
1991; Riley et al. 2003), Quebec (Mullins 1976; reported 
as dubious by Riley et al. 2003). U.S.A.: Arkansas (Riley 
et al. 2003), Connecticut (Britton 1920; Leng 1920; Wilcox 
1972; Mullins 1976; Downie and Arnett 1996; Riley et al. 
2003), District of Columbia (Riley et al. 2003), Kansas 
(Douglass 1929; Riley et al. 2003), Maine (Wilcox 1972, 
1975; Downie and Arnett 1996; Riley et al. 2003), Mary- 
land (Schroder et al. 1996; Riley et al. 2003), Michigan 
(Brown 1945; Wilcox 1972, 1975; Mullins 1976; Wheeler 
and Hoebeke 1979; Downie and Arnett 1996; Riley et al. 
2003), Missouri (Say 1826; Mullins 1976; Riley and Enns 
1979; Riley et al. 2003), New York (Wheeler and Hoebeke 
1979; Riley et al. 2003), North Carolina (Brimley 1938; 
Mullins 1976; Wheeler and Hoebeke 1979; Riley et al. 
2003), Ohio (Mullins 1976; Downie and Arnett 1996; 
Riley et al. 2003), Pennsylvania (Say 1826; Wilcox 1972, 


Vol. 119 


1975; Mullins 1976; Wheeler and Hoebeke 1979; Downie 
and Arnett 1996; Riley et al. 2003), Rhode Island (Sikes 
1999), Virginia (Mullins 1976; Wheeler and Hoebeke 1979; 
Riley et al. 2003), West Virginia (Clark 2000; Riley et al. 
2003). 

ILLINOIS: North Illinois, F. Knab Collection (1) 

Iowa: Johnson Co., lowa City, Wickham collector (1) 

WISCONSIN: Milwaukee Co., Milwaukee, Milwaukee 
Public Museum (1); Washburn Co., Spooner, 26 May 
1949, S. I. Parfin collector (1); Wisconsin, F. Knab 
Collection (1) 


Calligrapha sutureWa Schaeffer 1933 (Figure 1o) 

U.S.A.: New Hampshire (Schaeffer 1933). G6mez-Zurita et 
al. (2004) have used genetic data to support the elevation 
to specific status of this taxon, originally described as a 
local variety of C. mu/npunctata (Say 1824). Together with 
the re-description of the species, a number of new localities 
and new province and State records were provided which 
are reproduced below. 

MAniroBa: Franklin, 1649 feet, 17 July 2002, on Sa/zv 
Lebbiana, D. J. Funk collector (3) 

QueBEc: L’Amiante, Saint-Joseph-de-Coleraine, 27 
July 2000, on Sa/zx bebbiana, D. J. Funk collector 
(1); Lac-Saint-Charles, 26 July 2000, on Sa/ix eb- 
Diana, D. J. Funk collector (1); 

MAINE: Cumberland Co., Portland 18 June 2003, on 
Salix bebbiana, D. J. Funk (2); Kennebec Co., Sid- 
ney 6 June 2002, on Sa/ir Gebbiana, D. J. Funk col- 
lector (1); Kennebec Co., Sidney 20 June 2003, on 
Salix bebbiana, D. J. Funk collector (1); Somerset 
Co., Moose River 25 July 2000, on Sa/zx bebbiana, 
D. J. Funk collector (1) | 

MASSACHUSETTS: Berkshire Co., Hinsdale, 21 August | 
1898 (1); Hampden Co., Ludlow, 22 June 1902 (1); | 
Hampden Co., Wilbraham, J. O. Martin collector (1); 
Massachusetts (1) 

MICHIGAN: Baraga Co., 10 miles W Three Lakes, 7 
June 1982, David R. Smith collector (1); lower pen- 
insula, Ogemaw Co., Rt. I-75, 836 feet), 22 July 
2002, on Salix bebbiana, D. J. Funk collector (1); 
Agriculture Collection Michigan, 1891, C. F. B., F. 
Knab collector 1918 (1) 

MINNEsoTA: Aitkin Co., 3 miles S McGrath, 3 July 
1984, Downie and Wappes collector (1); Becker Co., 
Itasca St. Pk. Area, 28-29 June 1984, Downie and 
Wappes collector (1); Kanabec Co., Mora, 25 August 
1907, R. A. Vickery collector (2); St. Anthony Pk., | 
9 June 1907 (1) 

New HAmpsHIRE: Hillborough Co., Manchester, 16 
June, 25 June and 5 July 1932, W. S. Abbott collec- 
tor (4); Strafford Co., Milton, 3 July 1897, F. Knab | 
Collection 1918 (3); Squam Lake, F. Knab Collec- | 
tion 1918 (1) 

New York: Van Cortland Park, Charles Schaeffer coll. 
(1) | 

VERMONT: Windham Co., Brattleboro, spring 1894, | 
F. Knab Collection 1918 (1) | 

WISCONSIN: Wood Co., Cranmoor, 26 August 1909, - 
C. W. Hooker coll. (1) | 


2005 


Caliigrapha sylvia Stal 1860 (Figure 1o) 

U.S.A.: Arizona (Linell 1896; Wilcox 1972, 1975; Riley et al. 
2003), New Mexico (Townsend 1895; Riley et al. 2003). 
Mexico (Jacoby 1880-1888; Leng 1920; Linell 1896; 
Wilcox 1972, 1975; Blackwelder 1982). 

Note. This new record has to be considered provisional, since 
it seems to be from an intercepted specimen on the bor- 
der between Mexico and the US. However, the confirmed 
presence of the species in neighboring areas suggests that 
it could be naturally present in Texas too. 

TEXAS: El Paso Co., El Paso, 1 July 1942, with mus- 
tard greens from Mexico (1) 


Caliigrapha verrucosa (Suffrian 1858) (Figure |p) 

Canada: Alberta (Brown 1945; Robertson 1966; Bousquet 
1991; Riley et al. 2003), British Columbia (Brown 1945; 
Robertson 1966; Hatch 1971; Wilcox 1972, 1975; Bous- 
quet 1991; Riley et al. 2003), Manitoba (Brown 1945; 
Robertson 1966; Wilcox 1972, 1975; Bousquet 1991; Riley 
et al. 2003), Northwest Territories (Bousquet 1991; Riley 
et al. 2003), Ontario (Bousquet 1991; Riley et al. 2003), 
Saskatchewan (Brown 1945; Robertson 1966; Bousquet 
1991; Riley et al. 2003). U.S.A.: California (Riley et al. 
2003), Idaho (Hatch 1971; Riley et al. 2003), Montana 
(Leng 1920; Hatch 1971; Wilcox 1972, 1975; Riley et al. 
2003), Nebraska (Riley et al. 2003), Nevada (Riley et al. 
2003), Oregon (Hatch 1971; Wilcox 1972; Riley et al. 
2003), Utah (Riley et al. 2003), Washington (Hatch 1971; 
Riley et al. 2003), Wyoming (Lawson 1976; Riley et al. 
2003). 

ALASKA: Fairbanks North Star Co., Fairbanks, 16 July 
1952 (2); Fairbanks North Star Co., Fairbanks, 23 
May 1966, on Sa/zr spp., Joan Foote and Les Viereck 
collectors (2); Fort Yukon, 3 July 1953, R. I. Sailer 


(1) 


Caliigrapha vicina Schaeffer 1933 (Figure |p) 
Canada: New Brunswick (Bousquet 1991; Riley et al. 2003), 
Ontario (Brown 1945; Robertson 1966; Bousquet 1991; 
Riley et al. 2003), Quebec (Brown 1945; Robertson 1966; 
Bousquet 1991; Riley et al. 2003). U.S.A.: Maryland 
(Cavey 1994; Downie and Arnett 1996; Riley et al. 2003), 
New York (Schaeffer 1933; Brown 1945; Robertson 1966; 
Wilcox 1972, 1975; Downie and Arnett 1996; Riley et al. 
2003), Pennsylvania (Riley et al. 2003). 
CONNECTICUT: Connecticut, Blaisdell Collection (2) 
MASSACHUSETTS: Hampden Co., Montgomery, 21 
August 1897, F. Knab collector (1); Worcester Co., 
Southbridge, 3 May 1913, S. W. Bromley Collection 
(1) 
| OuI0: Ohio, C. F. B. collector (1) 
VERMONT: Bennington Co., Roberts Collection (1); 
Chittenden Co., Burlington, May 1942, U. V. M. 
collector (1) 


Zoogeography of Nearctic Coligrapha 

The catalogue of Cal//igrapha species in North 
America north of Mexico currently includes 38 taxa, 
| after establishing the synonymy for several subspecies 
pairs (Riley et al. 2001, 2003). The largest percentage 
(86.8%) corresponds to endemic forms for this part 


GOMEZ-ZURITA: NEW RECORDS AND BIOGEOGRAPHY OF C4ZLL/GRAPHA 95 


of the world, the only exceptions being species in a 
group that expands from Central America reaching 
the southernmost states in the United States (1.e., C. 
dislocata, C. multiguttata, C. serpentina, C. sylvia and 
C. fulvipes). The new distribution data presented here 
combined with the latest report on the distribution of 
the species in this genus (Riley et al. 2003) allows 
updating the data on endemicity of Ca//igrapha on a 
per country basis. The Canadian list of endemic Ca/- 
ligrapha includes the species C. amator, C. tiliae, and 
C. virginea, and the United States list the species C. 
amelia, C. androwt, C. cephalanti, C. floridana, and C. 
wickhami. The remaining species except for the above 
mentioned group expanding from Central America 
have been recorded in both countries. The highest 
diversity of Ca//igrapha can be found in the North- 
east quadrant of North America, in the area around 
the Great Lakes (Figure 2). 

An analysis of the ranges of distribution of all the 
North American species of Ca//igrapha helps estab- 
lishing a tentative separation of groups according to 
their geographical distribution (Table 1, Figure 2). 
This separation follows in general terms two gradi- 
ents: a longitudinal gradient most likely conditioned 
by the orography of the subcontinent, with the main 
mountain ranges following a North-South axis, and a 
latitudinal gradient possibly driven by climatic/eco- 
logical conditions. The first zonation distinguishes 
from west to east (1) a Pacific group limited eastward 
(less so to the north) by the Rocky Mountains, (11) an 
Atlantic group not surpassing this mountain system 
to the west, and (111) a Great Plains group distributed 
mainly in this geographical area, but spreading to the 
east and limited to the west by the Rockies (Table 1). 
The second possibly climatic or historical zonation 
includes (i) a Central American group, present only in 
the southernmost regions of North America, (1i) the 
endemic species from Florida, and (111) a very rich 
group in the area surrounding the Great Lakes (Table 
1). A group of a few trans-Nearctic species widely dis- 
tributed from coast to coast could be included among 
these too. Areas where two or more geographical 
assemblages overlap, on the edges from the centers 
of maximum diversity of each group, show a corre- 
sponding relative increase in species richness. This is 
well exemplified by the North-South corridor from 
Manitoba to Texas, where the Atlantic and Pacific 
groups meet intersecting with the Great Plains group, 
and showing richer Ca//igrapha faunas than the adja- 
cent regions (Figure 2). The observed distribution pat- 
tern of Ca//igrapha and the importance of the Cordil- 
leras (particularly the Rocky Mountains and Sierra 
Nevada) in separating the two dominant groups, Pacif- 
ic and Atlantic, is consistent with similar patterns ob- 
served for Onzychomys grasshopper mice, Gambelia 
lizards, Cvora/us rattlesnakes, A ge/enopsis spiders, and 
several plants, among others (e.g., Riddle and Honey- 
cut 1990; Orange et al. 1999; Pook et al. 2000; Hong, 
2001; Ayoub and Riechert 2004). The effective barrier 


96 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |. Grouping of Ca//igrapha species according to their distribution ranges in North America (north of Mexico). 


Longitudinal Latitudinal 
Pacific Great Plains _— Atlantic Trans-Nearctic Great Lakes Central American Florida 
C. sigmoidea C. incisa C. ignota C. californica C. alni C. dislocata C. cephalanti 
C. verrucosa C. praecelsis = C. knabi C. multiipunctata— C. suturella ~——C. multiguttata C. floridana 
C. rhoda C. rowena C. philadelphica C. alnicola C. serpentina 
C. scalaris C. bidenticola C. tiliae C. sylvia 
C. confluens — C. lunata C. vicina C. wickhamt 
C. spiraea C. virginea C. fulvipes 
C. amelia C. amator 
C. pnirsa C. apicalis 
€. androwi C. dolosa 
C. ostryae 
C. pruni 


of the Cordilleras could be not only responsible for 
the spatial structuring of the group in North America, 
but for its diversification as well. However, it is impor- 
tant to retain the idea that the effect of geographical 
barriers in the case of non-generalist phytophagous 
beetles like Ca//igrapha could be indirect, the primary 
effect being on the actual host plants of the insects. 

Latitudinal faunal structuring in North America, 
mainly related to climatic and the associated ecologi- 
cal gradients, has been analytically demonstrated for 
instance in mammals (Badgley and Fox 2000). At a 
different scale, a similar situation can be described for 
phytophagous specialist Ca//igrapha, where climate 
and ecology have a tight link through the distribution 
of the host plants. However, the observed latitudinal 
zonation in these beetles could be the effect of historical 
processes too, something that a phylogenetic analysis 
of genus can help to discern (G6mez-Zurita and coll- 
laborator, in preparation). The endemicity area of Flori- 
da could be related for instance to the prolonged insu- 
larization of this peninsula during the Pliocene, which 
has been proved effective to trigger the diversification 
of several animal groups (e.g., Gilbert 1987; Moler 
and Kezer 1993). 

The Great Lakes area is particularly interesting for 
two reasons: species diversity and species ecology. The 
highest species diversity of the genus in North America 
occurs in this region, particularly in the Canadian pro- 
vinces of Ontario and Quebec, resulting from the over- 
lap of the two richest species groups, the so-called 
“Great Lakes” and the Atlantic groups, together with 
the trans-Nearctic species. We believe that the high 
species diversity in this region is related to climatic 
and idiosyncratic features of this particular area that 
provides with a variety of niches for the colonization 
and possibly diversification of Ca//igrapha. An inter- 
esting feature of the North American Ca//igrapha sensu 
stricta is that they have exploited a trophic niche dif- 
ferent from the feeding selection of the other subgen- 
era and the congeneric species in Central and South 
America. While the latter feed on grasses and herba- 
ceous plants, most North American species feed on 


trees and shrubs from different botanical families, but 
typically found associated to streams and river banks 
(e.g., Alnus, Betula, Cornus, Ostrya, Physocarpus, 
Salix, Tilia, and U/mus, Brown 1945). Interestingly, 
these plants are particularly abundant and have the 
center of their distribution precisely in the Great Lakes 
area and North Eastern North America (Little 1980; 
USDA, NRCS 2004*), where most species of Ca//r- 
grapha are also found. This correlation is not surpris- 
ing and is particularly accurate in some cases. The 


ninebark, Physocarpus opulifolius (Rosaceae), is the | 
sole host-plant of C. spzraea, and the beetle has the | 
same as the core distribution of the plant in North | 


America (Figure In), which ranges from Quebec west 
to Michigan and south to Tennessee, although the plant 


reaches as isolated spots Eastern North Dakota and — 
Kansas to the west and Northern Florida to the north | 
(Wheeler and Hoebeke 1985; USDA, NRCS 2004*). | 
Tilia americana var. americana (American Basswood; © 
Tiliaceae) is restricted to the Northeastern quadrant of — 
North America (Little 1980; USDA, NRCS 2004*). | 
Three species of Ca//igrapha, including C. amator 
(Figure lc), feed on this host and are present only in © 
the plant center of distribution (Brown 1945). Similar 
situations can be described for C. 7gvora and their birch 

host Berula lenta (Betulaceae; possibly &. papyrifera | 
too), C. alni (Figure 1a), C. alnicola (Figure 1b), C. © 


amelia (Figure 1c), C. apica/is (Figure 1d), and C. con- 
Jluens (Figure 1g) on the alders A/zus incana americana 


and A. rugosa (Betulaceae), all living exclusively in 


North Eastern North America. C. sca/aris and U/mus 
americana (American Elm; Ulmaceae), and C. rhoda 
with Corylus americana (American Hazelnut; Betu- 


laceae), are all only present in the Eastern half of North | 
America. The trans-Nearctic Ca//igrapha (sensu stricta) 


species also conform to the expected correlated distri- 
bution with the host plant. So, Cornus sericea (= C. 
stolonifera, Dogwood; Cornaceae) and Salix bebbiana 
(Bebb’s Willow; Salicaceae), the preferred respective 
host plants of C. philadelphica and C. multipunctata, 


are also distributed throughout North America (Brown, | 


1945; USDA, NRCS 2004*). 


| 
| 
| 
| 


2005 


Leese : 


a [EP se, 


GOMEZ-ZURITA: NEW RECORDS AND BIOGEOGRAPHY OF C4ZL/GRAPHA 97 


_FiGure 2. Ca//igrapha species numbers in North American provinces and states. Thick lines roughly demarcate faunistic groups 


_ Calligrapha stands out among leaf beetles because 
it includes several parthenogenetic species, otherwise 
| very unusual among the Chrysomelidae (Brown 1945; 
| Robertson 1966; Cox 1996; Gémez-Zurita et al. 2004). 
It is noteworthy that all asexual species in Ca//igrapha 
| belong to the “Great Lakes” group and have typically 
\restricted ranges (with the possible exceptions of C. 
\alnicola and C. suturella, Figures 1b and 10). This 
|example of geographical parthenogenesis is however 
difficult to reconcile with current hypotheses relating 
\this reproductive mode with better colonizing abilities, 
highest adaptability through generalist behavior or 
jclonal microadaptation, and/or advantages on environ- 
ments with reduced biotic (particularly parasitic) inter- 


in the genus according to their geographic distribution. 


actions (Haag and Ebert 2004, and references there- 
in). The parthenogenetic Ca//igrapha species coexist 
in close sympatry with several other sexually reproduc- 
ing taxa (Brown 1945) and moreover we have hypoth- 
esized that interspecific hybridization might be the 
mechanism behind the origin of asexuality in Ca//r- 
grapha (Gé6mez-Zurita et al. 2004), not existing rea- 
sons a priori suggesting interspecific hybridization to 
occur following any geographical pattern. Again, phy- 
logenetic studies are needed to provide sound answers 
to these questions relevant for the biogeography, sys- 
tematics and evolution of such an remarkable genus, 
Calligrapha (G6mez-Zurita and collaborator, in prepa- 
ration). 


98 


Descriptive studies like this paper, aimed to resolve 
basic questions about the distribution of organisms in 
a particular geographical region, are critical to formulate 
meaningful hypotheses to be tested on a phylogenetic 
or ecological framework. The potential for information 
held at Museums and research institutions even for 
regions and taxa relatively well known, as for the leaf 
beetles from North America, is still enormous. A facil- 
itated access to these resources and ensuring conditions 
for their preservation prove very important to advance 
our understanding of the World’s biodiversity. 


Acknowledgments 

I express my gratitude to Alexander Konstantinov 
from the National Museum of Natural History (Smith- 
sonian Institution, Washington, DC) for the loan of the 
Calligrapha specimens for study. Thanks also to Daniel 
Duran, Charles and Susan Staines, and especially to 
Daniel Funk (Vanderbilt University, Tennessee) for col- 
lecting some of the specimens reported here. Thanks 
also to Anabela Cardoso for reading a preliminary ver- 
sion of this work and for her suggestions and ideas to 
improve it. Two anonymous referees provided very use- 
ful comments to further improve this work. 


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USDA, NRCS. 2004. The PLANTS Database, Version 3.5 
(http://plants.usda.gov). National Plant Data Center, Baton 
Rouge, Louisiana 70874-4490 USA. 


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Received 12 January 2004 
Accepted 7 March 2005 


_ their bills to carry pieces of suet weighing up to 33 


| (42-66 percent of bodyweight) from their bills to 


Weight-carrying Ability and Caching Behavior of Gray Jays, Fertsoreus 
canadensis. Adaptations to Boreal Winters 


LYNN L. ROGERS 
Wildlife Research Institute, 145 West Conan Street, Ely, Minnesota 55731 USA 


Rogers, Lynn L. 2005. Weight-carrying ability and caching behavior of Gray Jays, Perisoreus canadensis. Adaptations to boreal 
winters. Canadian Field-Naturalist 119(1): 101-104. 


During 16 August to 21 September 1984, I determined how Gray Jays (Perisoreus canadensis) carried flight-loads of different 
weights. Three individually identifiable Gray Jays weighing 60, 68, and 80 grams, used their bills to carry flight-loads weighing 
up to 33 percent of bodyweight but transferred heavier flight-loads from their bills to their feet 1-2 meters after takeoff. They 
had difficulty carrying flight-loads over 57 percent of bodyweight, and none attempted to carry flight-loads over 66 percent of 
bodyweight. By using their feet to bring heavy flight-loads closer to the center of lift, Gray Jays can carry heavier loads of meat, 
relative to body weight, than can Common Ravens (Corvus corax) which compete with Gray Jays at carcasses in winter and 
which do not carry objects with their feet. During 1969-2003, year-round observations near the southern edge of the Gray Jay 
range in northeastern Minnesota showed that caching behavior begins in August, continues over-winter, and ends at the onset 
of insect activity and green-up in early May. Gray Jays’ propensity to approach larger animals, including people, may not indicate 
unwariness but rather a superior ability and willingness to assess risks and food benefits. In the boreal forest in winter, risk of 
starvation is greater and risk of predation is lower than in relatively food-rich ecoregions farther south. 


Key Words: Ferisoreus canadensis, scavenging, food cache, flight-load, feet, competition, boreal forest, risk assessment, saliva, 
survival strategy, Minnesota. 


Gray Jay biology has been extensively researched, 
and their abilities to carry meat scraps in their feet and 
glue food caches to branches have been reported 
(Brewster 1937; Bock 1961; Dow 1965; Stirling 1968; 
Gill 1974; Bent 1946; Strickland and Ouellet 1993). I 
describe how Gray Jays carry flight-loads of different 
weights and report additional observations on scaveng- 
ing and caching behaviors in northeastern Minnesota 
near the southern edge of their range. 


Methods and Materials 

During 1969-2003, I made year-round observa- 
tions of Gray Jays feeding on suet or on White-tailed 
Deer (Odocoileus virginianus) that were killed by 
vehicles or Timber/Gray Wolves (Canis /upus) near 
Ely, Minnesota. During 16 August to 21 September 
1984, I determined weights of suet pieces carried off 
by three Gray Jays that were individually identifiable 
by their weights and by feather irregularities. I placed 
the suet pieces on a Hanson 500-g scale. When a Gray 
Jay landed and carried off one of the pieces (Figure 
1), I recorded the weight of the jay, the weight of the 
piece it selected, and how the jay carried it. 


Results 
Three Gray Jays, weighing 60, 68, and 80 g, used 


percent of bodyweight but transferred heavier loads 


their feet 1-2 meters after takeoff (Table 1). They had 
difficulty carrying flight-loads over 57 percent of 


bodyweight, and none attempted to carry pieces over —_ FicurE 1. The 500-gram scale used to weigh Gray Jays and 
66 percent of bodyweight. suet. Photo by Lynn Rogers. 


101 


102 


The Jays were successful in only 2 of 12 attempts to 
carry pieces of suet weighing 57-66 percent of body- 
weight (Table 1). In five attempts to fly with pieces 
weighing 58-64 percent of bodyweight, they dropped 
them while attempting to transfer them from their bills 
to their feet (Table 1). These dropped into deep grass 
and were abandoned. An 80-g Jay twice dropped a 46-g 
piece (58 percent of bodyweight) during transfers but 
on the third attempt reached an altitude of 2 meters and 
maintained level flight for over 50 meters. A 68-g Jay 
carried a 43-g piece of suet (63 percent of bodyweight) 
plus 2 g of stomach contents totaling 66 percent of 
bodyweight. It leaped into flight from a point 1.1 meter 
high, lost about a meter of altitude while transferring the 
load from its bill to its feet, and then flew close to the 
ground with labored flight. In three other instances, Jays 
used their bills to lift pieces of suet weighing 57-60 per- 
cent of bodyweight but did not attempt to fly off with 
them. 

With pieces of suet too heavy to fly off with, Jays 
held them with their feet, and used their bills to twist off 
small pieces. Amounts ingested per visit increased as 
the suet aged and softened. For example, the Jays in- 
gested an average of only 0.4 g per visit (n = 15 visits) 
from a fresh piece of hard suet on 19 August, but as the 
suet softened over the next 8 days, average amount 


TABLE |. Flight-loads carried by Gray Jays in their bills or feet in northeastern Minnesota, 16 August to 21 September 1984. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


ingested increased to 0.875 g per visit on 24 August 
(n = 8 visits), 1.45 g per visit on 25 August (n = 11 
visits), and 1.5 g on 27 August (n = 6 visits). Each Jay 
weighed 1-3 g more after the initial 1-3 visits, indicat- 
ing that they were retaining the initial 1-3 g for imme- 
diate digestion and flying off and storing what they 
ingested in subsequent visits. 

Gray Jays were common in the study area year- 
round. However, I observed no scavenging or caching 
of suet from the onset of green-up and insect activity in 
early May through the end of July. When caching be- 
havior resumed in August, they used their sticky saliva 
to glue caches to branches (Figure 2). 


Discussion 

With the exception of shrikes [Zanius spp.] (Sibley 
2001), I found no report of other passerines, including 
other corvids, carrying food or other objects with their 
feet. This ability to carry food with their feet aids com- 
petition for meat scraps at carcasses. By using their feet 
to bring flight-loads closer to their center of lift, Gray 
Jays can carry heavier loads, relative to bodyweight, 
than can Common Ravens, which do not carry food in 
their feet. For example, if ravens weigh 1200 g, as listed 
by Dunning (1984), an adult raven that carried a 237-g 
piece of meat to a tree carried only 20 percent of its 


Comments 


Flew easily. 

Flew easily. 

Flew easily carrying a Deer Mouse (Pevomyscus maniculatus). 
Jay rose 5 m at 45 degrees. 

Level flight but dropped food upon entering dense foliage. 


Ascended 60 degrees to fly over a 3-meter obstacle. 

Rose 3 m over a distance of 30 m. 

Flew easily. 

Flew easily. 

Level flight and glide. 

Lifted food with bill but did not attempt to fly with it. 

Dropped food 1-2 m after takeoff in first 2 attempts. Third attempt 


successful. 


Lifted food with bill but did not attempt to fly with it. 
Lifted food with bill but did not attempt to fly with it. 
Dropped food <1 m after takeoff during transfer from bill to feet 


and abandoned it. 


Level flight low to ground. 
Dropped food <2 m after takeoff during transfer from bill to feet 


and abandoned it. 


Bird Load 

weight weight —_ Percent of Means of 
(grams) (grams) bodyweight carrying 
60-80! 11 14-18% Bill 
80 14 18% Bill 
60-80! 14 18-23% Bill 
80 26 33% Bill 
80 26 33% Bill 
60 ZS 42% Feet 
60-80! 9 36-48% Beet 
60-80! 38 48-63% Feet 
68 36 53% Feet 
80 42 53% Feet 
80 43 54% Feet 
80 45 57% — 
80 46 58% Feet 
68 40 59% — 
68 4] 60% —- 
80 49 61% — 
68 43 63% * Feet 
68 43 63% _— 
80 51 64% _ 


Dropped food 1.5 m after takeoff during transfer from bill to feet 


and abandoned it. 


'Did not determine which of the three jays took the food, so the weight range of the three was used to calculate percentage of 


body weight. 


*This Jay ingested 2 g of suet in an earlier visit to the scale and still weighed 2 g heavier when carrying the 43 g, making the 


total load 45 g of suet—over 66% of body weight. 


2005 


ROGERS: WEIGHT-CARRYING AND CACHING BEHAVIOR OF GRAY JAYS 103 


FIGURE 2. A sunflower seed cached on the tip of a Red Pine (Pius resinosa) 
branch near Ely, Minnesota, in late September. Gray Jays use sticky 
saliva to encase food and glue it where it will remain above snow 
cover. It is unknown whether this encasement also preserves fresh- 
ness, protects against mold and bacteria, and/or deters insects and 
other cache-robbers by reducing odors, making caches distasteful, 
or disguising them as inedible items. Photo by Lynn Rogers. 


bodyweight, and an adult raven that dropped a 624-g 
piece of meat 1-2 meters after leaping into flight was 
unable to carry 52 percent of bodyweight (unpublished 
data). By comparison, Gray Jays fly buoyantly carrying 
that percentage of bodyweight in their feet (Table 1). 
Further, a 79.5-g male carried a 72.9-g piece of bread 
(92 percent of bodyweight) 20 meters in its feet before 
dropping it in Algonquin Provincial Park, Ontario 
(Strickland and Ouellet 1993). 

Gray Jays opportunistically scavenge ungulate car- 
casses that have been opened by carnivores, ravens, 
eagles, and other scavengers strong enough to penetrate 
tough skin. Gray Jays’ propensity to approach larger 
animals, including people, may not indicate unwariness 
but rather a superior willingness and ability to assess 
risks and food benefits. In the boreal forest in winter, 
risk of starvation is greater and risk of predation is 
lower than in relatively food-rich ecoregions farther 
south. For example, compared with the eastern decidu- 
ous forest, the boreal forest has longer winters, lower 
primary productivity, lower animal biomass (especially 
in winter when most birds have migrated), and lower 
diversity and abundance of predators (Odum 1971). 
Thus, selective pressures in the boreal forest, especially 
in winter, may favor more intensive assessment of 
potential food sources rather than the more cautious be- 
havior typical of migratory birds of the boreal forest 
| and of residents of ecoregions with more food and less 
| extreme winters. Gray Jays’ ability to carry heavy flight- 
| loads in their feet helps minimize exposure to predators 
| and competitors at carcasses. 


Gray Jays’ habit of using sticky saliva to encase their 
food caches and glue them to branches (Figure 2) 
enables them to place caches where they will remain 
above snow cover (Dow 1965). In northeastern Min- 
nesota, caching begins in August, leaving 3-4 months 
for spoilage and cache-robbing before winter. Tem- 
peratures in Ely, Minnesota, average 18°C in August, 
13°C in September, 6°C in October, and —-4°C in No- 
vember. Further study is needed to determine whether 
encasing caches in saliva preserves freshness, protects 
against mold and bacteria, and/or deters insects and 
other cache-robbers by reducing odors, making caches 
distasteful, or disguising them as inedible items. 

The winter diet is not entirely cached food and car- 
rion (Strickland and Ouellet 1993). On 26 January 
2002, a Gray Jay ingested fruticose lichens from the 
trunk of a Paper Birch tree (Bema papyrifera). | found 
no previous report of feeding on lichens, although eat- 
ing or caching fungi has been reported (Sutherland and 
Crawford 1979, Strickland and Ouellet 1993). 


Acknowledgments 
I thank A. J. Erskine, D. Strickland, and an anony- 
mous reviewer for helpful comments. 


Literature Cited 

Bent, A. C. 1946. Life histories of North American Jays, 
crows, and titmice. Smithsonian Institution United States 
National Museum Bulletin 191. United States Government 
Printing Office. [Reprinted by Dover Publications, Inc. 
1988. 495 pages, 68 plates]. 


104 


Bock, W. J. 1961. Salivary glands in the Gray Jays (Fev7- 
soreus). Auk 78: 355-365. 

Brewster, W. 1937. The birds of the Lake Umbagog Region 
of Maine. Bulletin of the Museum of Comparative Zoology 
66: 403-521. 

Dow, D. D. 1965. The role of saliva in food storage by the 
Gray Jay. Auk 82: 139-154. 

Dunning, J. B. 1984. Body weights of 686 species of North 
American birds. Monograph 1, Western Bird Banding 
Association. Eldon Publishing. P.O. Box 446, Cave Creek, 
Arizona 85331. 38 pages. 

Gill, D. 1974. The Gray Jay as a predator of small mammals. 
Canadian Field-Naturalist 88: 370-371. 

Odum, Eugene T. 1971. Fundamentals of ecology, 3" edition. 
W. B. Saunders Company. Philadelphia. 574 pages. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Sibley, David Allen. 2001. The Sibley guide to bird life and 
behavior. Chanticleer Press, Inc., New York, and Toronto. 
608 pages. 

Stirling, D. 1968. Notes on food and feeding habits of some 
wintering birds. Canadian Field-Naturalist 82: 14-17. 

Strickland, D., and H. Ouellet. 1993. Gray Jay (Perisoreus 

Canadensis). /n The Birds of North America (40). Edited 
by A. Poole, P. Stettenheim, and F. Gill. Academy of Natur- 
al Sciences, Philadelphia, and American Ornithologists’ 
Union, Washington, D.C. 

Sutherland, J. B., and R. L. Crawford. 1979. Gray Jay 

feeding on slime mold. Murrelet 60: 28. 


Received 13 September 2004 
Accepted 21 February 2005 


| 
| 
| 
7 


Premiéres mentions et répartition de la Salamandre sombre du Nord, 
Desmognathus fuscus, sur la rive nord du fleuve Saint-Laurent, au Québec 


JEAN-FRANCOIS DESROCHES! et DANIEL POULIOT? 


'Collége de Sherbrooke, Département des Techniques d’ écologie appliquée, 475 du Parc, Sherbrooke, Québec J1K 4K1 Canada 
Laboratoire d’écologie des eaux douces, Université du Québec a Trois-Riviéres, 3351 Boulevard des Forges, C.P. 500, Trois- 
Riviéres, Québec GOA 5H7 Canada 


Desroches, J.-F., et D. Pouliot. 2005. Premiéres mentions et répartition de la Salamandre sombre du Nord, Desmognathus Juscus, 
sur la rive nord du fleuve Saint-Laurent, au Québec. Canadian Field-Naturalist 119(1): 105-109. 


La Salamandre sombre du Nord (Desmognathus fuscus) est une espéce qui a fait l objet de plusieurs inventaires au Québec étant 
donné qu’elle y est a la limite nord de sa répartition. Nous rapportons les premiéres observations de cette espéce au nord du fleuve 
Saint-Laurent, depuis Trois-Riviéres, a 1’ ouest, jusqu’a Cap-Tourmente, vers l’est. Ces mentions ont été faites de 1987 a 2003, 
dans 20 sites. La majorité des sites ot fut découverte la Salamandre sombre du Nord sont des ruisseaux et des résurgences d’ eau. 


Mots-Clés: Desmognathus fuscus, Salamandre sombre du Nord, aire de distribution, répartition, habitat, Québec. 


The Northern Dusky Salamander (Desmmognathus fuscus) has been searched for extensively at its presumed northern range 
limit in Québec, south of the St-Lawrence River. We report the first records of this species north of the St-Lawrence River, 
from Trois-Riviéres, in the west, to Cap-Tourmente in the east. These observations were made between 1987 and 2003, at 
20 sites. Most of the sites where the Northern Dusky Salamander was found are streams and seepages. 


Key Words: Desmognathus fuscus, Northern Dusky Salamander, distribution, habitat, Québec. 


La Salamandre sombre du Nord (Desmognathus —_Un inventaire spécifique fut ainsi réalisé en 2002. II 
Juscus) se retrouve a la limite nordique de sa réparti- avait comme objectif de documenter la répartition de 
tion dans le sud de la province de Québec (Cook __l’espéce sur la rive nord du Saint-Laurent, et d’ obtenir 
1984). Les mentions historiques les plus nordiques de une idée générale de |’habitat de cette espéce a la 
lespéce ont été faites a Lac Trois-Saumons, comteé limite nordique de sa répartition. 
de I’ Islet (47°08'N, 70°12'O) et a Saint-André, comté 
de Kamouraska (47°38'N, 69°40'O), deux localités | Méthodologie 
situées sur la rive sud du fleuve Saint-Laurent (Den- L’inventaire de 2002 visant la découverte de la 
man 1963). Bleakney (1958) avait émis Il’ hypothese Salamandre sombre du Nord a d’ autres endroits sur la 
que le fleuve Saint-Laurent avait constitué une barriére rive nord du Saint-Laurent a été réalisé dans 40 nou- 


a la dispersion de la Salamandre sombre du Nord. veaux sites. Les sites ont été localisés préalablement 
Bider et Matte (1991), dans la version détaillée de sur des cartes topographiques 1: 50 000 ou choisis au 
l’Atlas des amphibiens et des reptiles du Québec, hasard sur le terrain. La méthodologie utilisée est 
mentionnent : «Puisque cette salamandre est active- similaire a celle de Weller (1977) et consiste en des 


ment recherchée par plusieurs herpétologistes, on peut recherches variant de 15 4 60 minutes dans les ruis- 
donc penser que sa répartition décrite ici est assez seaux et sur les rives de cours d’eau, en soulevant les 
complete». Or, en 1987, nous avons eu la surprise de —_ débris au sol. La présence et l’absence de la Sala- 
la découvrir pour la premiere fois sur la rive nord du mandre sombre du Nord, le nombre d’individus, leur 


Saint-Laurent stade de développement, |’ habitat, la date et la local- 

En 1987, JFD trouva quelques spécimens de |’e- isation au GPS en NAD83 ont été notés a chacun des 
speéce a Boischatel, a l’est de Québec, dans des eaux sites visités (annexe I). Les sites situés a proximité, 
de résurgence sur les rives abruptes de la riviére Mon- mais séparés par un secteur ot l’habitat est moins 
tmorency. Ils ont été formellement identifiés par lui- propice, ont été considérés et cartographiés séparé- 


méme en 1990, puis localisés approximativement dans ment malgré qu’ils puissent concerner le méme cours 
Atlas des amphibiens et reptiles du Québec (Bider et d’eau. Au moins un spécimen a été récolté a la plupart 
Matte 1994). Des recherches visant 4 découvrir d’ au- des sites, sauf lorsque ceux-ci étaient situés a prox- 
tres populations furent entreprises dés 1996 et 1998 imité l’un de l’autre. Les salamandres récoltées ont 
par les auteurs. En 2001, une Salamandre sombre dus été déposées au Musée canadien de la nature sous 
Nord fut trouvée par DP a la riviére Jean-Larose, prés les numéros CMNAR 35713-35729. 

du mont Sainte-Anne, 4 28 km au nord-est des men- 

tions faites a la rivigre Montmorency. Cette découverte Résultats et discussion 

ouvrait la porte a la possibilité que cette salamandre De 1987 4 2003, la présence de la Salamandre som- 
puisse étre plus répandue que prévu dans la région. bre du Nord a pu étre confirmée dans 20 sites situés 


105 


106 


sur la rive nord du Saint-Laurent. Cing d’entre eux 
ont été découverts de 1987 a 2001, 13 durant I’ inven- 
taire spécifique de 2002, et deux durant des recherch- 
es supplémentaires effectuées en 2003 plus a |’ ouest. 
Ces 20 sites s’étalent d’ouest en est de la riviere 
Saint-Maurice, a Trois-Rivieres, jusqu’a Cap-Tour- 
mente (figure 1). Les sites situés aux extrémités (ouest 
et est) sont séparés par 160 km. Toutes les observa- 
tions ont été faites dans les contreforts du Bouclier 
Canadien, dans le domaine de I’ Erabliére a Tilleul, ot 
l’ altitude moyenne varie de 55 a 116 m (Robitaille et 
Saucier 1998). Au sein de cette aire, la Salamandre 
sombre du Nord semble en général localisée a des 
secteurs ot le microhabitat est propice: eaux de suin- 
tement, zones vaseuses avec mousses par endroits et 
bordure des ruisseaux. Le long des grandes rivieres 
ou on la retrouve, comme les riviéres Saint-Maurice, 
Batiscan et Montmorency, elle fut retrouvée dans 
des eaux de ruissellement localisées et habituellement 
situées a quelque distance de la riviére (de quelques 
metres a plus de 300 m), et non de facon réguliere sur 
les rives des cours d’eau. 

Les habitats dans lesquels fut trouvée la Salamandre 
sombre du Nord sont : les ruisseaux (< 3m de largeur) 
(59,0 % des sites), les eaux de résurgence (36,4 %), 
et un fossé forestier sans eau (4,6 %). Ces habitats 
concordent avec ceux décrits dans la littérature : habi- 
tats forestiers aquatiques Ou semi aquatiques, souvent 
abondante aux endroits ou se retrouvent des résur- 
gences d’eau et le long des petits ruisseaux forestiers 
bordés de roches ou autres abris (Bishop 1941; Pet- 
ranka 1998). 

La plus forte proportion de ruisseaux peut représen- 
ter un biais. En effet, l’?espéce semble assez typique 
des eaux de résurgence, mais ces milieux sont diffi- 
cilement repérables sur des cartes a l’échelle 1: 50 000, 
comparativement aux cours d’eau permanents qui sont 
bien visibles. 

Au Canada, la Salamandre sombre du Nord est une 
espéce associée aux Appalaches (Bleakney 1958). Nos 
résultats démontrent cependant qu’elle semble bien 
établie sur la rive nord du Saint-Laurent, dans les 
contreforts du Bouclier Canadien. Comment |’ espéce 
s’y est-elle rendue? A quel endroit a-t-elle traversé le 
fleuve Saint-Laurent? La topographie du territoire 
porte a croire que ce fut dans le secteur de la ville de 
Québec, et vraisemblablement a partir de l’embou- 
chure de la riviére Chaudiére (rive sud du Saint- 
Laurent, en bordure du pont de la route 73) qu’elle 
aurait franchi le fleuve, tel que proposé par Bider et 
Matte (1994). La Salamandre sombre du Nord se re- 
trouve dans la riviere Chaudiére et a pu traverser le 
fleuve a cet endroit, ot la largeur de ce dernier est 
plus étroite (moins de | km) qu’aux autres secteurs 
(2 km et plus). De plus, a cet endroit, les Appalaches 
se rendent a proximité du Saint-Laurent, ce qui n’est 
pas le cas plus a l’ouest ou une large bande de la val- 
lée du Saint-Laurent sépare les Appalaches du fleuve. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Suite a son arrivée sur la rive nord. la Salamandre 
sombre du Nord se serait ensuite répandue vers |’ ouest, 
dans le Bouclier Canadien, pour se répandre jusque 
dans la région de Trois-Riviéres, quelques 115 km plus 
a l’ouest. Vers l’est, il semble que sa répartition s’ arréte 
aux environs du Cap-Tourmente, ot |’ altitude devient 
plus importante et la topographie plus accidentée. La 
végétation y change également, passant du domaine de 
l’Erabliére a Tilleul 4 celui de la Sapiniére 4 Bouleau 
jaune (Robitaille et Saucier 1998). D’ ailleurs, ce secteur 
constitue la limite de répartition nord-est de plusieurs 
especes végétales et animales, dont certains amphi- 
biens et reptiles (Cook 1984). On ignore si la Sala- 
mandre sombre du Nord s’est répandue vers le nord 
mais les recherches effectuées ces derniéres années 
n’ont pas permis de la repérer. Le secteur situé a I’ ouest 
de Trois-Riviéres, situé lui aussi dans le domaine de 
l’Erabliére 4 Tilleul, semble particuligrement propice, 
mais a cet endroit la vallée du Saint-Laurent est pré- 
sente sur une bande plus large et le Bouclier Cana- 
dien se retrouve donc plus au nord. Nos recherches 
effectuées plus a l’ouest dans le Bouclier Canadien, 
dans les Laurentides au nord de Montréal et en Outa- 
ouais, n’ont pas permis de trouver de Salamandres 
sombres du Nord mais seulement la Salamandre a 
deux lignes (Zurycea bislineata). 


Perspectives 

La découverte de la Salamandre sombre du Nord 
sur la rive nord du Saint-Laurent ouvre la porte a la 
possibilité que certaines espéces d’amphibiens, notam- 
ment les salamandres, soient répandues plus largement 
que l’on croit au Québec, et notamment sur la rive 
nord. En effet, la présence de cette espéce sur la rive 
nord du Saint-Laurent était ignorée jusqu’en 1990 
(année ou les premiers spécimens furent identifiés), 
malgré le fait que les salamandres de ruisseaux (la 
Salamandre sombre du Nord, la Salamandre a deux 
lignes et la Salamandre pourpre (Gyrinophilus por- 
Phyriticus) aient fait P objet de quelques inventaires 
professionnels au Québec (Bider et Matte 1991; Pen- 
dlebury 1973; Bleakney 1958). Il apparait possible 
que la Salamandre sombre du Nord soit encore plus 
répandue sur la rive nord du Saint-Laurent, de méme 
que sur la rive sud vers l’est. Des inventaires spéc- 
ifiques réalisés dans le futur pourraient permettre de 
tester cette hypothese. 


Remerciements 

Les auteurs remercient tous ceux qui ont participé 
aux travaux de terrain : Daniel Banville, Josiane 
Bergeron, Joél Bonin, Nathalie Coté, Benoit Couture, 
Frédérick Desroches, Sébastien Desroches, Ian-Erik 
Gosselin, Isabelle Picard, Martin Savard et Vicky 
Tremblay. Ils remercient également Frederick W. Sch- 
ueler, pour ses commentaires sur la version préliminaire 
du texte, de méme que Mario Darsigny pour la réal- 
isation de la carte de I’ aire d’étude et de la localisation 
des sites. 


2005 DESROCHES ET POULIOT: LA SALAMANDRE SOMBRE AU QUEBEC 107 


72200' O 


4 


kilometres 
Echelle: 1: 600 000 


FiGuRE |. Localisation de |’aire d’étude au Québec (en médaillon) et des 20 sites ot fut trouvée la Salamandre sombre du 
Nord sur la rive nord du Saint-Laurent, de 1987 a 2003. 
1. Trois-Riviéres, MRC de Francheville. 46°23'35"N; 72°38'58"0. 
2  Saint-Narcisse, MRC de Francheville. 46°33'14"N; 72°24'59"0. 
3. Municipalité de paroisse de Saint-Alban, MRC de Portneuf. 46°46'14"N; 72°09'50"0. 
4. Municipalité de paroisse de Saint-Alban, MRC de Portneuf. 46°46'04"N; 72°09'25"0. 
5. Municipalité de Cap-Santé, MRC de Portneuf. 46°41'36"N; 71°50'33"0. 
6. Municipalité de Cap-Santé, MRC de Portneuf. 46°41'17"N; 71°49'18"0. 
7. Municipalité de paroisse de Saint-Augustin-de-Desmaures, MRC de la Communauté Urbaine de Québec. 
46°47'19"N; 71°31'08"0. 
8. Charlesbourg, MRC de la Communauté Urbaine de Québec. 46°55'35"N; 71°14'54"0. 
9. Charlesbourg, MRC de la Communauté Urbaine de Québec. 46°54'44"N; 71°16'00"0. 
10. Boischatel, MRC de la Cote-de-Beaupré. 46°53'45"N; 71°09'07"0. 
11. Boischatel, MRC de la Céte-de-Beaupré. 46°53'25"N; 71°08'47"0. 
12. Sainte-Pétronille, MRC de I’Ile-d’ Orléans. 46°51'10"N; 71°06'55"0. 
13. Sainte-Famille, MRC de I’[le-d’ Orléans. 46°57'49"N; 70°58'24"0. 
14. Sainte-Famille, MRC de I’Ile-d’ Orléans. 46°57'54"N; 70°58'17"0. 
15. Saint-Francois, MRC de 1’Ile-d’ Orléans. 47°01'12"; 70°50'16"0O. 
16. Mont-Sainte-Anne, MRC de la-Céte-de-Beaupré. 47°05'04"N; 70°54'01"0. 
17. Saint-Joachim, MRC de la-Céte-de-Beaupré. 47°04'13"N; 70°51'03"0. 
18. Cap-Tourmente, MRC de la-Céte-de-Beaupré. 47°04'55"N; 70°47'09"0. 
19. Cap-Tourmente, MRC de la-Céte-de-Beaupré. 47°04'59"N 70°46'42"0. 
20. Cap-Tourmente, MRC de la-Céte-de-Beaupré. 47°05'05"N; 70°46'16"0. 
Lesroutes provincials 20, 40, 55, 73, et 573 sont inclues. 


Littérature citée Bishop, S. C. 1941. The salamanders of New York. New 
Bider, J.-R., ct S. Matte. (compilé par). 1991. Atlas des York State Museum Bulletin (324). Albany, New York. 
| amphibiens et reptiles du Québec 1988-1989-1990, version 365 pages. 
détaillée. Société d’histoire naturelle de la vallée du Saint-  Bleakney, J. S. 1958. A zoogeographical study of the am- 
Laurent et ministére du Loisir, de la Chasse et de la Péche phibians and reptiles of Eastern Canada. National Museum 
du Québec. Québec. 429 pages. of Canada, Bulletin 155. 119 pages. 
| Bider, J.-R., ct S. Matte. 1994. Atlas des amphibiens et Cook, F. R. 1984. Introduction aux amphibiens et reptiles 
reptiles du Québec. Société d’ histoire naturelle de la val- du Canada. Musée national des sciences naturelles et 
lée du Saint-Laurent et ministére de l’Environnement et Musées nationaux du Canada. Ottawa, Canada. 211 


de la Faune du Québec. Québec. 106 pages. pages. 


108 THE CANADIAN FIELD-NATURALIST Vol. 119 


Denman, N. S. 1963. A range extension of the Dusky Sa- _— Petranka, J. W. 1998. Salamanders of the United States 


lamander in Quebec. Canadian Field-Naturalist 77 :62. and Canada. Smithsonian Institution Press. Washington 
Pendlebury, G. B. 1973. Distribution of the Dusky Sala- and London. 587 pages. 

mander Desmognathus fuscus fuscus (Caudata: Pletho- Robitaille, A., et J.-P. Saucier. 1998. Paysages régionaux 

dontidae) in Quebec, with special reference to a pop- du Québec méridional. Les Publications du Québec. 213 

ulation from St. Hilaire. Canadian Field-Naturalist 87: pages + carte. 

131-136. Weller, W. F. 1977. Distribution of stream salamanders in 
southwestern Quebec. Canadian Field-Naturalist 91 : 
299-303. 


Received 16 February 2004 
Accepted 17 February 2005 


Annexe I. Détails sur les localités et les salamandres observées 


Chaque donnée est présentée de la maniére suivante : # du site (voir la figure 1), nom de la municipalité, nom de la MRC 
(municipalité régionale de comté), localisation GPS en NAD 83, type d’habitat (eaux de résurgence, ruisseau) et présence 
ou non de Salamandre a deux lignes, date, nombre d’individus de Salamandres sombres du Nord observés et stade, nombre 
de spécimens récoltés, nom des observateurs, numéro de catalogue musée canadien de la nature (amphibiens et reptiles) 
CMNAR. 


Iie 


Les Vieilles-Forges, MRC de Francheville. 46°23'35"N; 72°38'58"O. Environ 100 m a l’ouest de la riviére Saint- 
Maurice. Habitat : Eaux de résurgence. 
20 juillet 2003 : 2 adultes (1 récolté), obs. : J.-F. Desroches, D. Pouliot, I. Picard et N. Coté. CMNAR 35278. 


Saint-Narcisse, MRC de Francheville. 46°33'14"N; 72°24'59"0. Parc de la riviére Batiscan, environ 300 m a |’ ouest de 
la riviére Batiscan. Habitat : Eaux de résurgence. 

20 mai 2003 : 1 subadulte, obs. : D. Pouliot et M. Savard. 

24 mai 2003 : 4 adultes (1 récolté) et 2 subadultes. : D. Pouliot et N. Coté. CMNAR 35729. 


Municipalité de paroisse de Saint-Alban, MRC de Portneuf. 46°46'14"N; 72°09'50"O. Sime rang, a 1,5 km au sud du 
lac en Coeur. Habitat : Eaux de résurgence. Présence de Salamandres a deux lignes. 
20 aoat 2002 : 5 adultes (1 récolté), JFD-02.10, obs. : D. Pouliot. CMNAR 35722. 


Municipalité de paroisse de Saint-Alban, MRC de Portneuf. 46°46’04’’N; 72°09’25’’O. 5®™° rang, a 1,5 km au sud du 
lac en Coeur. Habitat : ruisseau. 
20 aott 2002 : 2 adultes, obs. : D. Pouliot. 


Municipalité de Cap-Santé, MRC de Portneuf. 46°41'36"N; 71°50'33"0. 2,3 km a l’est de Portneuf, au nord de la route 
138. Habitat : ruisseau. Présence de Salamandres a deux lignes. 
29 juillet 2002 : 1 adulte (récolté) et 1 jeune, JFD-02.17, obs. : D. Pouliot. CMNAR 35725. 


Municipalité de Cap-Santé, MRC de Portneuf. 46°41'17"N; 71°49'18"0. Ruisseau André-Jacques, 2,8 km au nord- 
ouest de Cap-Santé, entre |’ autoroute 40 et la route 138. Habitat : ruisseau. Présence de Salamandres a deux lignes. 
22 juillet 2002 : 5 adultes (1 récolté) et | jeune (récolté), JFD-02.19, obs. : D. Pouliot. CMNAR 35727. 


Municipalité de paroisse de Saint-Augustin-de-Desmaures, MRC de la Communauté Urbaine de Québec. 46°47'19"N; 
71°31'08"0. 1,3 km a lest de la route 367. Habitat : ruisseau. 
9 aout 2002 : 4 adultes (1 récolté), JFD-02.18, obs. : D. Pouliot. CMNAR 35726. 


Charlesbourg, MRC de la Communauté Urbaine de Québec. 46°55'35"N; 71°14'54"0. Secteur du lac Des Roches. 
Habitat : fossé forestier. 
31 mai 1998 : | adulte, obs. : J.-F. Desroches et F. Desroches. 


Charlesbourg, MRC de la Communauté Urbaine de Québec. 46°54'44"N; 71°16'00"0. Riviere Des Roches, en aval du 
lac Des Roches. Habitat : ruisseau. Présence de Salamandres a deux lignes. 

17 mai 1996 : 3 adultes, obs. : J.-F. Desroches. 

9 juillet 1997 : 1 subadulte, obs. : J.-F. Desroches. 

31 juillet 2002 : 1 adulte (récolté) et 1 jeune, JFD-02.16, obs. : D. Pouliot. CMNAR 35724. 


Boischatel, MRC de la C6te-de-Beaupré. 46°53'45"N; 71°09'07"0. Riviére Montmorency en amont des chutes. Habitat : 
eaux de résurgence et ruisseau. Présence de Salamandres a deux lignes. 

1987 : plusieurs trouvées; 2 adultes (JFD-70) et 1 larve (JFD-55) récoltés, obs. : J.-F. Desroches. CMNAR 35715, 35714. 
23 avril 1990 : 2 adultes et quelques larves, obs. : J.-F.Desroches. 

13 octobre 1990 : 2 adultes, obs. : J.-F. Desroches. 

28 avril 1991 : 2 adultes, obs. : J.-F. Desroches. 

30 mai 1991 : 35 adultes et jeunes + 10 larves, obs. : J.-F. Desroches. 

17 avril 1992 : 1 adulte et 1 larve, obs. : J.-F. Desroches. 

9 février 1993 : | larve, obs. : J.-F. Desroches. 


2005 DESROCHES ET POULIOT: LA SALAMANDRE SOMBRE AU QUEBEC 109 


1S. 


14 


iS. 


16. 


18. 


19. 


20. 


20 mars 1994 : | larve, obs. : J.-F. Desroches et F. Desroches. 

3 mai 1995 : 2 adultes, obs. : J.-F. Desroches. 

14 mars 1996 : 6 adultes (3 récoltés) et 2 larves, JFD-385, obs. : J.-F. Desroches et B. Couture. CMNAR 35717. 
8 mai 1996 : 5 adultes et 3 jeunes, obs. : J.-F. Desroches. 

17 mai 1997 : 5 adultes et 3 larves, obs. : J.-F. Desroches. 

12 septembre 1999 : 2 adultes et 2 jeunes, obs. : J.-F. Desroches, J. Bonin et I. Picard. 

18 octobre 1999 : 2 adultes, 1 jeune et | larve, obs. : J.-F. Desroches. 

28 aoit 2002 : 4 adultes et | jeune, obs. : D. Pouliot et D. Banville. 


. Boischatel, MRC de la Cote-de-Beaupré. 46°53'25"N; 71°08'47"0. Riviere Montmorency au pied des chutes. Habitat : 


eaux de résurgence. Présence de Salamandres a deux lignes. 

1990 : quelques trouvées, obs. : J.-F. Desroches. 

8 juillet 1991 : 32 adultes et jeunes, obs. : J.-F. Desroches. 

12 septembre 1999 : 2 adultes et 1 sub-adulte, obs. : J.-F. Desroches, J. Bonin et I. Picard. 
18 octobre 1999 : 2 larves, obs. : J.-F. Desroches. 


. Sainte-Pétronille, MRC de I’ fle-d’ Orléans. 46°51'10"N; 71°06'55"O. Ruisseau Marie-Anne, secteur ouest de I’lle d’ Orléans, 


200 m au nord de la route 368. Habitat : ruisseau. Présence de Salamandres a deux lignes. 
14 juin 2002 : 5 adultes et 2 jeunes, obs. : D. Pouliot, V. Tremblay, D. Banville et N. Cété. 


Sainte-Famille, MRC de I’fle-d’ Orléans. 46°57'49"N; 70°58'24"O. Secteur centre de I’Ile-d’ Orléans, 1,2 km au sud-ouest 
de Sainte-Famille, au nord de la route 368. Habitat : ruisseau. Présence de Salamandres a deux lignes. 
14 juin 2002 : | adulte, obs. : D. Pouliot, V. Tremblay et N. Coté. 


Sainte-Famille, MRC de I’ Ile-d’ Orléans. 46°57'54"N; 70°58'17"O. Habitat : ruisseau. Présence de Salamandres a deux 
lignes. 
25 juillet 2002 : 1 adulte, JFD-02.15, obs. : D. Pouliot. CMNAR 35723. 


Saint-Francois, MRC de I’[le-d’Orléans. 47°01'12"; 70°50'16"O. Ruisseau du Moulin, secteur est de I’Ile-d’ Orléans, 
800 m au nord de la route 368. Habitat : ruisseau. Présence de Salamandres a deux lignes. 
17 juin 2002 : 2 adultes, obs. : D. Pouliot et N. Coté. 


Mont-Sainte-Anne, MRC de la-Céte-de-Beaupré. 47°05'04"N; 70°54'01"O. Riviere Jean-Larose, 840 m au nord de la route 
360. Habitat : eaux de résurgence. Présence de Salamandres a deux lignes. 

15 aodt 2001 :1 adulte (récolté), JFD-01.10, obs. : D. Pouliot et I.-E. Gosselin. CMNAR 35718. 

10 juin 2002 : 1 adulte (photographié), obs. : D. Pouliot, V. Tremblay et D. Banville. 


. Saint-Joachim, MRC de la-Céte-de-Beaupré. 47°04'13"N; 70°51'03"0. Ruisseau des Carriéres, en bordure est de l’av- 


enue Royale. Habitat : eaux de résurgence. Présence de Salamandres a deux lignes dans ruisseau a proximité. 
4 juillet 2002 : 1 adulte, JFD-580, obs. : J.-F. Desroches et V. Tremblay. CMNAR 35719. 


Cap-Tourmente, MRC de la-Céte-de-Beaupré. 47°04'55"N; 70°47'09"0. Ruisseau de la Friponne. Habitat : eaux de 
résurgence et ruisseau. Présence de Salamandres a deux lignes. 
28 aodt 2002 : 2 adultes (1 récolté), JFDA-02.9, obs. : D. Pouliot, D. Banville et V. Tremblay. CMNAR 35721. 


Cap-Tourmente, MRC de la-Céte-de-Beaupré. 47°04'59"N 70°46'42"0. Ruisseau Pierré. Habitat : ruisseau. Présence 
de Salamandres a deux lignes. 
28 aout 2002 : 2 adultes, obs. : D. Pouliot, D. Banville et V. Tremblay. 


Cap-Tourmente, MRC de la-Céte-de-Beaupré. 47°05'05"N; 70°46'16"0. Ruisseau Pierré. Habitat : ruisseau. Présence 
de Salamandres a deux lignes. 
28 aout 2002 : 2 adultes (1 récolté), JFD-02.8, obs. : D. Pouliot, D. Banville et V. Tremblay. CMNAR 35720. 


First Records of Long-beaked Common Dolphins, De/phinus capensis, 
in Canadian Waters 


JOHN K. B. FORD 


Fisheries and Oceans Canada, Pacific Biological Station, 3190 Hammond Bay Road, Nanaimo, British Columbia V9T 6N7 
Canada 


Ford, John K. B. 2005. First Records of Long-beaked Common Dolphins, De/phinus capensis, in Canadian Waters. Canadian Field- 
Naturalist 119(1): 110-113. 


The genus De/phinus has recently been determined to be comprised of two species, the Short-beaked Common Dolphin, D. 
delphis, and the Long-beaked Common Dolphin, D. capensis. D. delphis is regularly observed in eastern Canadian waters, but 
is known only from a single stranding in British Columbia. Two specimen records and a series of sightings of D. capensis in 
British Columbian waters during 1993-2003, detailed here, are the first for this species in Canada. D. capensis normally ranges 
only as far north as central California, and its abundance in those waters increases in association with warm-water oceanographic 
events. Although the species appears to be rare in British Columbia, future sightings during warm-water periods might be 
anticipated. 


Key Words: Long-beaked Common Dolphin, De/phinus capensis, Short-beaked Common Dolphin, De/phinus delphis, stranding, 


entanglement, feeding. 


Dolphins of the genus De/phinus have a wide but 
disjunct distribution in warm temperate to tropical 
waters worldwide from about 40-60°N to roughly 
50°S (Jefferson et al. 1993). Considerable uncertainty 
about the taxonomic status of De/phinus species has 
long existed, especially in the eastern North Pacific 
(Banks and Brownell 1969; van Bree and Purves 1972; 
Heyning and Perrin 1994). Dall (1873) proposed a 
new species De/phinus bairdi Dall from coastal waters 
of California, which he considered distinct in rostral 
length and colouration from De/phinus delphis Lin- 
naeus, previously known from the Atlantic Ocean. 
The validity of D. dairdii was challenged by some 
authors (e.g., True 1889; van Bree and Purves 1972) 
and supported by others (e.g., Miller 1936; Banks and 
Brownell 1969). Until this uncertainty was resolved 
recently, the names PD. bairdii, D. delphis bairdii, and 
D. delphis were applied inconsistently to eastern North 
Pacific De/phinus (Guiguet 1954; Norris and Prescott 
1961; Heyning and Perrin (1994) recently reviewed 
the taxonomic status of De/phinus and provided clear 
evidence for two distinct species in the northeastern 
Pacific, the Short-beaked Common Dolphin, D. de/- 
Phis Linnaeus, and the Long-beaked Common Dol- 
phin, 2. capensis Gray. D. capensis is equivalent to 
the original long-beaked form described as D. bairdii 
Dall, but the latter name is now considered a junior 
synonym and is not used (Rice 1998). Short-beaked 
and Long-beaked Common Dolphins can be distin- 
guished reliably by colouration pattern and the propor- 
tional length of the rostrum in adults (ratio of rostral 
length to zygomatic width in D. delphis is 1.21-1.47 
and in D. capensis is 1.52-1.77; Heyning and Perrin 
1994). The validity of these two species has been cor- 
roborated by genetic analyses (Rosel et al. 1994) and 


110 


they are now widely accepted (Rice 1998; Carretta 
et al. 2002; Reeves et al. 2002). 

Short-beaked and Long-beaked Common Dolphins 
have generally allopatric ranges in warm-temperate and 
tropical regions of the Pacific and Atlantic Oceans, 
though they are narrowly sympatric in some coastal 
waters (Heyning and Perrin 1994; Perrin 2002). D. 
capensis appears to prefer shallower and warmer water 
than D. de/phis, and is most often found within 50 nm 
of shore. In the northeastern Pacific, 2. de/phis ranges 
along the coast and pelagically from South America 
to southern British Columbia, while 2. capensis is 
found primarily in the Gulf of California and along 
the west coast of Baja California north to central Cali- 
fornia (approximately 36°N; Carretta et al. 2002, Per- 
rin 2002). 

The status of De/phinus in Canadian waters was 
reviewed by Gaskin (1992) before D. capensis was 
recognized as distinct from D. de/phis. Common Dol- 
phins are frequently seen during summer and fall on 
the banks offshore of Nova Scotia, and occasionally 
range as far north as coastal waters off Newfoundland 
(approximately 50°N; Gaskin 1992). The species in 


this region is D. delphis, as D. capensis has not been 7 


recorded north of 20°N in the western Atlantic (Per- 


rin 2002). In western Canadian waters, the genus has f} 


until recently been represented only by a single indi- — 


vidual found stranded on southern Vancouver Island | 


in April 1953. This specimen, an adult male, was des- 
cribed by Guiguet (1954) as D. dairdi7, but the animal J 
was judged to be D. de/phis by Heyning and Perrin | 

(1994) based on colouration evident in a photograph 
in the 1954 article (W. Perrin, National Marine Fish- 
eries Service, La Jolla, California, personal commu- | 
nication). To confirm this identification, the skull of | 


2005 ForD: LONG-BEAKED COMMON DOLPHINS IN CANADIAN WATERS 


the specimen was measured and found to have a ros- 
tral length to zygomatic width ratio of 1.37, clearly 
within the range diagnostic of D. de/phis (Heyning 
and Perrin 1994). (Royal British Columbia Museum, 
Victoria, British Columbia.; catalogue number 5792, 
examined 12 January 2004). The following thus des- 
cribes the first records of Long-beaked Common Dol- 
phins, D. capensis, in Canadian waters. 

The earliest record of D. capensis in British Colum- 
bia is from 2 February 1993, when a male Common 
Dolphin stranded alive at Mayne Island and died shortly 
thereafter (Table |). Post mortem examination at the 
provincial Animal Health Centre revealed the animal to 
be in poor condition, with extensive osteomyelitis of 
the caudal vertebrae and a heavy parasitic infection. The 
animal was recorded as DP. de/phis, but no skeletal 
material or measurements were collected (R. Lewis, 
Animal Health Centre, Abbotsford, British Columbia, 
personal communication) Archived photographs of the 
specimen were subsequently located and examined, 
which showed the colouration pattern distinctive of D. 
capensis. 

The species is next recorded in a series of sight- 
ings in the late summer and fall of 2002. On 21 
August 2002, a single individual was observed and 
photographed in Port McNeill, northeastern Vancou- 
ver Island, British Columbia. The animal appeared 
lethargic and in poor health. During the period of 18 
September to 2 October 2002, a pair of D. capensis 
was sighted on several occasions in Vancouver har- 
bour. One of these individuals, shown in Figure 1, 
was determined from dorsal fin markings to be the 
same animal as in the earlier Port McNeill sighting. 
Next, a group of four Common Dolphins was ob- 
served and video recorded on | November 2002 at 
Goose Bay, on the central coast of British Columbia 
} north of Vancouver Island. Examination of the video 
recording showed the animals to be D. capensis. 

The last set of records involves a pair of D. capensis 
| photographed in Victoria harbour, southern Vancouver 
Island, on 19 September 2003. In one of these animals, 
| the distal 5-8 cm portion of the rostrum was bent at an 
| approximately 45° angle to the left. This distinctive 
| individual had been sighted previously, together with 
} another Long-beaked Common Dolphin (probably the 
| Same companion as in Victoria), on several occasions 
| approximately 150 km to the south, in Puget Sound, 
Washington, during July-August 2003 (S. Norman, 
National Marine Fisheries Service, Seattle, Washington, 
) personal communication). On 7 October 2003, a pair of 
\dolphins, including the animal with the deformed 
|Tostrum, became entangled in a gillnet during a salmon 
jtest fishery in Alberni Inlet, on the west coast of Van- 
couver Island. The animal with the deformed rostrum 
was released alive from the net, but the other animal 
died and was collected and frozen for later examination. 
On 28 October 2003, the released dolphin was again 
sighted near the location of the 7 October entaglement. 


TABLE |. Records of Dedphinus capensis in British Columbian waters. All species identifications were made from photographs or video recordings of the animals’ distinctive coloura- 


tion, except for the entangled animal collected on 7 October 2003, the identity of which was confirmed by cranial measurements.. 


Number Comments 


Type 


Position 


Location 


Date 


Stranded alive, died en route to rehab facility. 


Animal appeared lethargic 


Stranding 
Sighting 


48°S1'N, 123°18'W 
50°35'N, 127°05"W 


49°16'N, 123°8'W 


Mayne Island 


2 February 1993 
21 August 2002 


Port McNeill 


Series of sightings of apparently same pair; one 


Sighting 


Vancouver harbour 


18 September — 2 October 2002 


individual same as 21 August 2002 sighting from 


dorsal fin markings 


Tight group of 4 individuals swimming actively 


in bay; observed for approx. 2 hours. 
Same pair as 7 October 2003 


One died, one released 


Sighting 


31°22'N, 127° 39°W 


Goose Bay 


| November 2002 


Sighting 


123° 23°W 
49°16'N, 123°8"W 
49°16'N, 123°8'W 


Victoria harbour 48°25'N, 


Alberni Inlet 


19 September 2003 
7 October 2003 


Entanglement 
Sighting 


Same animal released on 7 October 2003 


Alberni Inlet 


28 October 2003 


112 


se, 


THE CANADIAN FIELD-NATURALIST 


Volhahe 


ies 


FiGurRE |. One of a pair of De/phinus capensis observed in Vancouver harbour, 19 September 2003, showing the distinctive 
colouration of the species (see Heyning and Perrin (1994) or Reeves et al. (2002) for comparative illustrations of D. 


capensis and D. delphis). Photo by Anne Carson. 


Post mortem examination of the entangled animal 
showed that it had been in good health and had been 
feeding. It was a male, 187 cm long and 80 kg in 
weight. Tooth rake scars on its body were positively 
matched to scars in photographs taken on 19 Septem- 
ber 2003 of the animal accompanying the dolphin 
with the deformed rostrum. Its stomach contained 
remains of four Pacific hake (Werluccius productus) 
and six Pacific herring (C/upea pallasi). The skeleton 
of the dolphin was prepared and is now part of the 
collection of the Natural History Museum, Malaspina 
University-College, Nanaimo, British Columbia (cat- 
alogue number M-0283). The ratio of rostral length 
to zygomatic width is 1.69, well within the range 
measured for D. capensis (1.52-1.77; Heyning and 
Perrin 1994). 


These recent records confirm that D. capensis 


occurs in British Columbian waters, but it is unlikely 
more than a rare visitor. The records presented here 
are the northern-most for the species in the eastern 
North Pacific. Previously, there had been no sighting 
or stranding records for 2. capensis north of central 


California, though the species has only recently been | 


differentiated from PD. de/phis in sighting surveys 


(Heyning and Perrin 1994; Carretta et al. 2002). D. 
delphis is common in both inshore and offshore waters _ 


off California, but no confirmed sightings have been 


made in waters off Oregon and Washington (Carretta _ 
et al. 2002). Of four stranding records of Common — 


Dolphins on the coasts of Oregon and Washington, 


only one, a D. de/phi's, was identified to species (Car- — 


retta et al. 2002). There are no confirmed records of 


D. delphis in British Columbia since the 1953 strand- | 
ing. In Californian waters, the abundance of the two © 
species varies both seasonally and inter-annually, with | 
highest densities associated with warm-water events — 
(Heyning and Perrin 1994, Forney and Barlow 1998). | 


Heyning and Perrin (1994) noted that the proportion 


of D. capensis versus D. delphis strandings on the coast | 
of California increased for several years following the — 
warm-water El Nifio of 1982-83. The appearance of D. | 
capensis in British Columbian waters may be associat- — 


ed with increased water temperatures, and future sight- 


ings might be anticipated during warm-water periods. — 


2005 


Acknowledgments 

I thank the following for their assistance: W. Perrin, 
J. Heyning and R. Pitman for confirming species iden- 
tification from photographs; G. Ellis, L. Barrett- 
Lennard, and M. McDonald for information on sight- 
ings; R. Lewis, A. Carson and E. Westle for providing 
photographs; M. O and L. Spaven for collecting the 
entangled specimen, J. Watson for facilitating its post 
mortem examination, and S. Raverty for undertaking 
the necropsy; S. Crockford for identification of stomach 
contents; L. Kennes for providing access to the skull 
of the 1953 specimen at the Royal British Columbia 
Museum; S. Norman for information on De/phinus 
sightings in Washington state; and D. Nagorsen for 
helpful comments on an earlier draft of the manu- 
script. 


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FORD: LONG-BEAKED COMMON DOLPHINS IN CANADIAN WATERS 113 


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Received 3 August 2004 
Accepted 7 March 2005 


Occurrence, Composition and Formation of Auppia, Widgeon Grass, 
balls in Saskatchewan Lakes 


RANDY W. OLSson!, JosEF K. ScHMuTz?, and U. THEODORE HAMMER* 


'W. P. Fraser Herbarium, University of Saskatchewan, 51 Campus Drive, Saskatoon, Saskatchewan S7N 5A8 Canada 

*Centre for Studies in Agriculture, Law and Environment, University of Saskatchewan, 51 Campus Drive, Saskatoon, 
Saskatchewan S7N 5A8 Canada 

3Corresponding author Schmutz. joe.schmutz @usask.ca 

* Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada 


Olson, Randy W., Josef K. Schmutz, and U. Theodore Hammer. 2005. Occurrence, composition and formation of Ruppia, 
Widgeon Grass, balls in Saskatchewan Lakes. Canadian Field-Naturalist 119(1): 114-117. 


Widgeon Grass (Ruppia maritima) is an aquatic vascular plant (Ruppiaceae) which has been the source for rare balls of plant 
material found at the shores of lakes on four continents. In North America, the lakes involved were in North Dakota, Oregon, 
and now northern and southern Saskatchewan. The formation of the balls has not been observed in nature, but similar balls 
have been produced in other studies with Aosidonia or Turtle Grass (Hydrocharitaceae) fibers under the wavelike action in a 
washing machine. Our samples are from a saline lake in southern Saskatchewan (49°N), and an over 40-year-old sample from 
an unknown lake north of the boreal transition zone (52°N). Comparisons of the plant material with herbarium specimens 
confirm that the balls are almost entirely comprised of Auppia maritima, with minor items including invertebrate animal parts, 


sand pebbles and feathers. The context in which the material was found is consistent with the proposition that they are formed _ 


by Ruppia inflorescences breaking apart, drifting to near shore due to wind and being rolled into balls by wave action. 


Key Words: Ruppia maritima, Widgeon Grass, plant balls, saline lake, Saskatchewan. 


Unusual balls of plant material were encountered at —_ dry in some or most years. On 20 October 2000 there 
shores of two different lakes in northern and southern —_ was no standing water covering the white salt flat. 


Saskatchewan. Judging from available literature, similar A second set of plant balls was found during lake | 
balls have been found in Europe, North America and studies in the 1950s by D. S. Rawson (e.g., Rawson and | 
New Zealand (Cannon 1979; Essig 1948; Gerbeaux and Moore 1944). The balls were given to UTH about | 
Ward 1986). While the actual formation of these balls 1960, but data on time and location of collection were _ 
has not been observed in nature, the combined action _ not recorded. The collection has been given to Glenn | 
of waves on fragments of aquatic vegetation near shores is — Sutter, for inclusion in the natural history collections | 
thought to be involved. Such balls have, however, been —_ of the Royal Saskatchewan Museum in Regina (RSM | 


produced using /osidonia or Turtle Grass (Hydro- Accession 17443). 
charitaceae) fibers under the wave-like actions simulat- 
ed by a washing machine (Cannon 1979). Results and Discussion 


Names for the plant balls exist in Dutch, English, More than 50 plant balls were first found on 6 , 
French, German and Italian, 15 names in total (McAtee July 2001 by JKS along the southeastern shore of | 


1925). Some of these names date back to classical ttmes | Sandoff Lake (Figure 1) where the prevailing winds 
(E. A. Parry 77 Cannon 1979). This attests toa wide- —_ are northwesterly. The balls were scattered in a band 
spread, even if rare, occurrence of these balls. The = approximately 3 m wide and 100 m long, at the high 


names also attest to a degree of fascination; a curiosity | water mark where the pebbly shore merges with short- 
likely enhanced by the unpredictable occurrence of __ grass prairie. The balls were still present, although more | 


these structures in nature. encrusted in salt, on 30 June 2003. Their position in a 
single layer, similarity in salt or algal coverage and 
Sample Sources slightly compressed form suggests that only one genera- 


Our collection includes six balls of plant material in _ tion of such balls occurred during the three years. Their | 


two sets. One set was located at Sandoff Lake (49°00'N, _ position at the upper reaches of the shore suggests that 
104°18'W) in the Lake Alma Upland within the Mis- __ the balls may have been washed there during strong | 
souri Coteau of south-central Saskatchewan. Sandoff — winds but not returned to the lake with the receding | 
Lake is a | km’ saline lake in a hill-and-dale landscape. _ water during calmer weather. A walk around the lake re- | 
Sandoff Lake has no above-ground outflow, and lies in vealed two sets of balls on the east but none on the west 

the Missouri-Mississippi River drainage (Fung et al. (primarily upwind) shore. | 
1999). As is typical of many saline lakes, it is probably Analysis of the material in the two sets of balls | 
fed through ground water in addition to runoff and — showed that they were composed mainly (greater than | 
springs visible on the lake’s shore. Sandoff Lake goes 95%) of Ruppia inflorescence stems, peduncles, and | 


114 


2005 


fruits. The fruits were black, ca. 2 mm long, and beaked. 
These identifications were based on comparisons be- 
tween the samples and specimens from the W. P. Fraser 
Herbarium, and descriptions from several floras (e.g., 
Haynes 2000). In three of the balls from Sandoff Lake, 
Ruppia maritima L. was identifiable. Small amounts of 
other material found included twigs, small pebbles, and 
a feather. 

Saskatchewan has two species of Ruppia. R. cirrhosa 
(Petagna) Grande (synonym = &. occidentalis S. Wat.) 
and R. maritima L. (Harms 2003). Ruppia maritima \s 
found in the southern third of Saskatchewan, and Aup- 
pia cirrhosa\s found in the southern half of the province 
(Hammer and Heseltine 1988; W. P. Fraser Herbarium). 
In North America, Ruppia maritima is found throughout 
eastern and western coastal regions (Haynes 2000). 
Ruppia cirrhosa\s found from Alaska south to Texas 
and From California east to the Great Lakes (Haynes 
2000). 

The two species of Awppia belong to Ruppiaceae, the 
Ditch-grass family, also known as Widgeon Grass 
(Haynes 2000), The one genus includes approximately 
10 species worldwide, and two in North America. Aup- 
pia is an annual, or rarely perennial, sessile herb sub- 
mersed in brackish or saline water. Flowering occurs 
spring-fall. 

The plant balls as shown by Essig (1948), and 
Swanson and Springer (1972) are highly similar to our 
material. Also, ®. maritima L. seems to be the primary 
plant species involved, at least in North America (Table 
1). This suggests that the conditions that favour ball for- 
mation are relatively specific. The involvement of saline 


OLSON, SCHMUTZ, AND HAMMER: AUP?P/A BALLS IN SASKATCHEWAN 


LS 


FIGURE |. Ruppia maritima balls located at the southwest 
shore of Sandoff Lake in Saskatchewan, on 6 July 
2001. 


FIGURE 2. Two Ruppia maritima balls. One (left, 6.0 cm diametre) from Sandoff Lake, and 
the larger (right, 13.5 cm) from northern Saskatchewan. 


116 THE CANADIAN FIELD-NATURALIST Vol. 119 
TABLE |. Characteristics of plant balls as reported in the literature and found in our Saskatchewan samples. 

Site Name Location Lake Type Plant Species Reference 
No name NE North Dakota Saline Ruppia maritima SS 1972 
Mediteranean - Sea shore Posidonia oceanica JFMC 1979 
No name Near Dawson, ND Saline Ruppia maritima FPM 

Miller Lake Oregon — “similar” to FPM GWF 

Little Borax Lake California Saline Ruppia maritima EOE 1948 
Sandoff Lake S Saskatchewan Saline Ruppia maritima This report 
Unknown N Saskatchewan Unknown Ruppia maritima This report 


References: SS = Swanson and Springer 1972, JEMC = Cannon 1979, FPM = EP. Metcalf 7 McAtee 1925, GWF=G. W. | 


Field zz McAtee 1925, EOE = E.O. Essig 1948. 


lakes may not be crucial to ball formation but merely 
reflect that Ruppia is a saltwater species. The stems of 
Ruppia are delicate, relying on the surrounding water 
to support their mass (Kantrud 1991*). The stems are 
likely susceptible to breakage during periods of unusual 
turbulence. The stem bases also decay as the plants 
reach senescence (Kantrud 1991*). These two factors 
would leave large amounts of plant material in the water 
in late summer and/or fall, likely leading to the homoge- 
neous content of the Auppia balls. 

Given the structure of the balls, the location in which 
they were found and explanations from the literature, 
we suggest that the following mechanisms are involved 
in their formation. The fibrous stalks of Auwppia are 
prone to breakage and to sticking together. Two balls 
that were cut (Figure 2) and one that was teased apart 
suggest that the material 1s Rwppra throughout and does 
not contain any other material in its core. When the 
stems of the Appia break and tangle together, floating 
mats accumulate and are driven by wind to the shore 
(e.g., Essig 1948). Here, the repeated wave action in 
combination with the resistance provided by the shore 
in shallow water rolls the mat back and forth. This 
rolling action was thought responsible for producing 
balls of plant material in the lint trap of a washing 
machine (Cannon 1979). Finally, on windy days the 
balls may be washed up onto shore out of reach of the 
lapping waves on calm days that might otherwise return 
the balls to water where they may decay. The distribu- 
tion of balls at Borax Lake (Essig 1948) suggests that 
the balls were deposited over time as water evaporated 
and the shallow lake receded, differing slightly from our 
accumulation at the shore. Judging from the misaligned 
fibers in our material, interconnected in seemingly all 
directions (Figure 2), we doubt that the balls have 
grown in layers (e.g., Essig 1948). Instead, the different 
sized balls may simply be due to different amounts of 
plant material in a mat that starts to roll on the shore. In 
the Sandoff Lake sample, salt spray had encrusted the 
balls and, combined with grass shoots growing through 
the balls, had firmly anchored the balls on the upper 
shore. The Sandoff Lake balls tended to be oval. We 
suggest that they were round originally but that the 
material settled to an oval shape after they were covered 
in salt, dried and became anchored on the shore. 


The formation of balls by wave/wind action and 
friction is not limited to aquatic plants. Near Blaine 
Lake, Saskatchewan, wind combined with large and 
wet snowflakes led to snowball formation observed 
on the packed snow in a yard (Julie Hupé, personal 


communication). Pierce et al. (2004) compared amor- | 
phous, decomposed remains of large animals that | 
washed up on a Chilean shore in 2003 (Chilean Blob), | 
with other such “carcasses” variously reported as sea | 
monsters for over a century. Such sightings occurred in | 
2), West Coast of Tasmania, and 
Northeast Coast of the United States in addition to | 
Chile. The authors identified the material as decom- | 


Bermuda (n = 


posed remains of large whales. The material was held 


together by cross-hatched layers of collagen fibers. | 


This flexible layering of collagen in a blob was differ- 
ent from the firm arrangement of the inflorescence 
stems in our Avppra balls. 


Acknowledgments 


Dennis Dyck kindly took the photo of the balls and | 
Blaine Novakowski sliced them for analysis. Glenn | 
Sutter cataloged the material as part of the Royal Sask- | 
atchewan Museum’s collections (RSM Accession | 


17443). 


Documents Cited (marked * in text) | 

Kantrud, H. A. 1991. Wigeongrass (Ruppia maritima L.): | 
A literature review. U.S. Fish and Wildlife Service, Fish | 
and Wildlife Research 10. Jamestown, North Dakota: | 
Northern Prairie Wildlife Research Center Home Page. 
http://www.npwrc.usgs.gov/resource/literatr/ruppia/rup 
pia.htm (Version 16JUL97). 


Literature Cited 

Cannon, J. F. M. 1979. An experimental investigation of osv- 
donia balls. Aquatic Botany 6: 407-410. | 

Essig, E. O. 1948. The Auppia balls of Little Borax Lake. | 
Science Monthly 66: 467-471 | 

Fung, K., B. Barry, and M. Wilson. Z7zors. 1999. Atlas of 
Saskatchewan. University of Saskatchewan, Saskatoon, | 
Saskatchewan. 

Gerbeaux, P., and J. Ward. 1986. The disappearance of | 
macrophytes and its importance in the management of | 
shallow lakes in New Zealand. Pages 119-124 7 Proceed- | 
ings of the 7" International Symposium on Aquatic Weeds, | 
European Weed Research Society and Association of | 


2005 


Applied Biologists. Loughborough University of Tech- 
nology, Loughborough, Leicestershire, England, 15-19 
September. 

Hammer, U. T., and J. M. Heseltine. 1988. Aquatic macro- 
phytes in saline lakes of the Canadian prairies. Hydrobi- 
ologie 158: 101-116. 

Harms, V. L. 2003. Checklist of the vascular plants of Saskat- 
chewan, and the provincially and nationally rare native 
plants in Saskatchewan: Including important synonyms, 
authorities, common names, and various status indicators. 
University of Saskatchewan Extension Press, Saskatoon, 
Saskatchewan. 

Haynes, R. R. 2000. Ruppiaceae Hutchinson, Ditch-grass 
Family. Pages 75-76 77 Flora of North America. Edited by 
the Flora of North America Editorial Committee, Volume 
22, Oxford University Press, New York. 


OLSON, SCHMUTZ, AND HAMMER: AUPP/A BALLS IN SASKATCHEWAN 


ey 


McAtee, W. L. 1925. Notes on drift, vegetable balls, and 
aquatic insects as a food product of inland waters. Ecology 
6: 288-302. 

Pierce, S. K., S. E. Massey, N. E. Curtis, G. N. Smith, Jr., 
C. Olavarria, and T. K. Maugel. 2004. Microscopic, 
biochemical and molecular analysis of the Chilean Blob and 
a comparison with the remains of other sea monsters: Noth- 
ing but whales. Biological Bulletin 206:125-133. 

Rawson, D.S., and J. E. Moore. 1944. The saline lakes of 
Saskatchewan. Canadian Journal of Research 22: 141-201. 

Swanson, G. A., and P. F. Springer. 1972. Widgeongrass balls 
on alkali lakes. The Prairie Naturalist 4(2): 52-54. 


Received 23 April 2004 
Accepted 11 February 2005 


Why and How to Study a Snowcover 


WILLIAM O. PRUITT, JR. 


Taiga Biological Station and Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada 


Pruitt, William O., Jr. 2005. Why and how to study a snowcover. Canadian Field-Naturalist 119(1): 118-128. 


Specialized terminology, duration, thickness, hardness, density of the snowcover of taiga and tundra are described. Methods 
for detailed study of these characteristics are given, with description and use of simple as well as specialized instruments and 
techniques in relation to winter ecology of mammals and birds. 


Key Words: snow, snowcover, taiga, tundra, snow ecology, winter ecology, boreal ecology, snow instruments, snow terminology. 


Why is it necessary to study and record the details of 
a snowcover? Is not knowledge of the thickness of the 
cover (or even just its presence or absence) sufficient to 
explain the use or lack of use of an area by animals? In 
this paper I will show how details of a snowcover affect 
not only the presence or absence of a species but also 
how a species can use the snowcover, how it is affected 
by the variations in the snowcover morphology or 
sometimes is induced to emigrate from a region because 
of characteristics of only one layer in the snowcover 

Moreover, some species move down from the surface 
of the snowcover or up into the snowcover itself for 
varying periods of time. For some species the snowcov- 
er acts as a hindrance to travel or to obtaining food from 
under it, to others as a blanket protecting life from the 
deep cold of Full Winter (Pruitt 1957, 1960). This paper 
will also describe and review widely-accepted, as well 
as specialized, descriptors, instruments and measuring 
techniques. These techniques give one the tools for 
basic descriptive analyses of snow features. Sturm 
(1992) demonstrated, based on such a framework, an 
analysis of heat flow through taiga api as affected by 
qali fall and bare qamaniq. 


Terminology 

The English language evolved in a misty, maritime 
climate where snow was an uncommon occurrence, 
consequently it is woefully deficient in words represent- 
ing snow phenomena. The “scientific” languages (Latin 
and Greek) are derived from cultures which had even 
less familiarity with snow and its various forms than did 
English. Therefore, it seems best to use precise words 
that designate features and concepts from cultures that 
have evolved closer ecological links with snow than our 
Euro-Canadian one. Familiarity with these words ex- 
poses us to a whole new world of snow. The participants 
in the Scandinavian-Canadian Field Workshop on 
Rangifer-Snow Ecology (Pruitt 1971) resolved “...to 
assemble all known snow terms in all northern lan- 
guages and to illustrate each term with a photograph or 
accurate drawing. We believe that publication of such 
a lexicon would materially advance the study of boreal 
ecology. We invite submission of snow terms and pho- 


118 


tographs for possible inclusion in the lexicon.” Table 1 
consists of some words from such cultures that my stu- 
dents and I have found especially useful in our studies 
of snow ecology (Pruitt 1978, 1979, 1999). The simpli- 
fied spelling and pronunciation of these words is given, 
enclosed in brackets, immediately after the first use in 
this paper. 

There are four major qualities of a snowcover, 
particularly as they affect living organisms: Duration, 
Thickness, Hardness, and Density (Pruitt 1984a). The 
latter two characteristics enter into calculations of 
“snow water equivalent” and are influenced primarily 
by wind and the occurrence of winter thaws or freeze- 
thaw cycles. Thus there are four combinations that gen- 
erally agree with four major geographic types of snow- 
cover: 
steppes and coastal regions with freeze-thaw and wind, 
tundra with wind and no freeze-thaw, 
inland southern regions with freeze-thaw but no wind, 
taiga with no freeze-thaw and no wind. 


This classification has recently been confirmed in gen- | 


eral by Sturm et al (1995). Confusion sometimes still — 


exists between “arctic” and “‘taiga” snow processes, | 


especially concerning the known effects of snowcover 


on large mammals such as wolves, foxes and caribou | 


(Olsson et al. 2003). 


Duration 

The onset and disappearance of a snowcover, 
whether taiga api [ah-PEE] or tundra upsik [OOP-sik] 
(Table 1), are accompanied almost always by fluctua- 
tions in snowcover over the landscape. Inexperienced 
researchers have a tendency to fail to record the details 
of exact position and shape of the transitory spots where 
api first forms and where it lingers longest. One should 
make an effort to record these characteristics, preferably 
by detailed sketches including measurements of the 


spots and their exact locations, as well as times they 


occur. These spots and connecting areas will probably 
be where the Hiemal Threshold (Pruitt 1957) will occur 
first. The sequence of Fall Thermal Overturn, Fall Criti- 


cal Period, Hiemal Threshold, Full Winter, Hiemal Ter- | 


mination, Spring Critical Period, Spring Thermal Over- 


2005 PRUITT: WHY AND How To STUDY A SNOWCOVER 119 


turn (Pruitt 1978, Figure 4-1; 1984b) are all important 
events in the cycle of the seasons for the plants and ani- 
mals of the forest floor. The duration and intensity of the 
fall and spring critical periods can govern survival of 
populations of small mammals (Whitney and Feist 
1984). For some individuals or age-classes of a popula- 
tion of Caribou (Rangifer tarandus), overwinter survival 
seems to be a race between decreasing body mass 
(“loss of condition’) and duration of the snowcover. 

One of the important environmental factors affecting 
snow is wind. If wind is lacking, the complex, six- 
armed snow crystals drift down and settle undisturbed 
on other crystals which have preceded them, or onto the 
ground and short vegetation. Here, over time, they will 
undergo metamorphosis driven by the heat and mois- 
ture rising from the earth below (Schemenauer et al. 
1981; Seligman 1936). This heat and moisture will 
modify the snow crystals by subliming water molecules 
from the attenuated tips of the arms and redepositing 
them on colder crystals farther from the earth. The 
bottom-most crystals are the oldest and, therefore, have 
undergone this process the longest. Consequently, they 
are the most modified. Sometimes they are completely 
eroded away and their molecules have been redeposited 
higher up in the snow cover. Because of the physics of 
snow crystals the modified ones are in the shape of six- 
sided scrolls or pyramids. 

They can reach 10 mm or more in size and, upon 
magnification, they look as if they are made from tiny 
glass logs (Figure 1). The metamorphosed layer can 
extend 10 cm or more up into the cover and is properly 
known as pukak [POO-kak]. This layer is of great 
importance to subnivean mammals and winter-active 
invertebrates. In some subalpine or northern taiga 
regions where there is intense heat-loss from the snow- 
cover surface, almost all the snowcover can consist of 
pukak. 

Such is the idealized situation, with the api affected 
only by the heat and moisture rising from the earth, and 
with little or no heat coming from the supranivean air. 
In temperate and maritime climates where warm, moist 
air masses can intrude on the winter climate there can 
be heat and moisture gradients moving downwards as 
well. In these conditions pukak may not form, or may 
be severely modified (Pruitt 1984b). On the island of 
Newfoundland, with its relatively warm, wet, yet 
snowy, maritime climate, I found pukak in only a few 
sites near the upper limit of trees in the Long Range 
Mountains. I suspect that the general lack of pukak is 
as reasonable an explanation for the island’s markedly 
depauperate small mammal fauna as is any species’ 
inability to cross the Strait of Belle Isle (Pruitt 1968). 
Indeed, I now believe that pukak, not only its physical 


..} condition as a safe winter habitat for small mammals 


and invertebrate life, but as an indicator of general win- 
ter conditions for all boreal life deserves a great deal 
more research attention than it has had in the past. 


TABLE |. Specialized snow terminology of some northern peo- 
ples. Transliterations in parentheses after aboriginal word indi- 


cate pronunciations in general English sounds. (Capitalized 


syllable means stress accent; 


indicates a hard, glottal stop) 


English Kobuk Valley (Alaska) 
Inupiat 
Snow anniu (ah-NEE-u) 


Snow on trees 


Dense crystals, moist air 
hitting very cold surface 


Snow on the ground,soft 
fluffy taiga snow 


Base of snowcover, with 
large, pyramidal crystals 


Hard, wind-beaten snow 
(tundra or prairie) 


Drifting snow 


qali (KAH-lee) 
kanik (KAH-nik) 
api (ah-PEE) 
pukak (POO-kak) 
upsik (OOP-sik) 


siqog (see-KOK) 


Smooth snow surface of salumaroaq 
very fine particles (sah-loo-MAH-roak) 
Rough snow surface of natatgonaq 


large particles 


(nah-tat-GO-nak) 


Sun crust siqoqtoaq 
(see-KOK-toe-ak) 
Drift downwind of an kimoaqruk 


obstruction 


(kee-mo-AK-rook) 


Space between drift and anyemanya 
obstruction causing it (ahn-ye-MAHN-ya) 
Arrowhead-shaped drift kalutoganik 

moving over upsik (kah-lu-toe-GAHN-ik) 
Wind-eroded upsik kaioglaq 

from kalutoganik (kai-OHG-lak) 
Irregular surface caused tumarinyiq 


by differential erosion 


(too-mar-IN-yik) 


Bowl-shaped depression qamaniq 

in api around base of tree (KAH-mahn-ik) 
Dindye (Fort 
Yukon, Alaska) 

Snowcover thick enough detthlo(k) 

to need snowshoes (DET-thlo(k)) 
Russian 

Spot blown bare of snow vyduvi 
(vih-DOO-Vv1) 

Area of thick snow cover, zaboi 

persists all summer (ZAA-boy) 
Finnish 

Animal’s overnight burrow kieppi 

in the snow cover (KEY-eppi) 

Kanik crystals, vertical Huurre 

surface (HOO-rreh) 

Kanik crystals, Kurra 

horizontal surface (KU-rra) 


120 


In spring, especially in alpine regions and in the High 
Arctic, the long hours of intense solar radiation can melt 
or sublime the snowcover surface and meltwater can 
trickle down through the upsik and refreeze against the 
ground (Miller and Kiliaan 1980). This phenomenon is 
known to Sami reindeer herders as Cuokki [CHU-ok-ki] 
(Eriksson 1976; Pruitt 1979). Miller and Kiliaan (1980) 
outlined the severe effects of Canadian High Arctic 
cuokki on large mammals such as Peary Caribou 
(Rangifer tarandus pearyi). 


Thickness 

Thickness is governed by latitude, proximity and 
direction from major sources of atmospheric moisture, 
as well as the regional climate. Thickness, especially in 
mountainous regions, can affect supranivean animals 
simply by hiding or “smoothing over” portions of the 
overwintering habitat. In steep, rocky or irregular terrain 
this can result in falls causing broken limbs. Murie 
(1935) noted the importance of snowcover in the migra- 
tions and seasonal distribution of Alaska Caribou, most 
of which are essentially mountain animals. Edwards 
and Ritcey (1959) discussed the effects of snowcover 
thickness on the altitudinal migrations of Moose (A/ces 
alces) in British Columbia and Edwards (1956) corre- 
lated snowcover thickness and trends in ungulate popu- 
lations. One of the classic publications in snow ecology 
is Nasimovich (1955) who demonstrated the influence 
of snowcover thickness on wintering aggregations of 
Moose in the Ural Mountains. 

Mammals as large as Fisher (Martes pennanti) occa- 
sionally burrow into the api in a kieppi [KEY-eppi], 
as a hunting tactic or to escape extreme cold. Tundra 
mammals such as Arctic Fox (A/opex lagopus), Red 
Fox (Vulpes vulpes) or even Wolverine (Gulo gulo) 
will seek shelter by burrowing into soft spots in upsik 
or inside the curl-space of a snow cornice or a drift 
(Pruitt, unpublished observations. ). 

Grouse-like birds frequently use kieppi to escape 
low temperatures and predators (Formozov 1970; Kor- 
honen 1980a, 1980b; Marjakangas 1986). The latter 
worker noted that four major factors have to be con- 
sidered when defining the influence of api conditions 
on construction of kieppi: (1) the presence of crusts 
within the api, (2) the penetrability of these crusts, 
(3) the thickness of any soft snow on top of the crust, 
and (4) the total api thickness. Penetrability mainly 
depends on hardness and thickness of the crust. Crusts 
of up to 2000 to 3000 gm cm? (20 to 30 N) vertical 
hardness are “readily penetrable,;’ whereas harder 
crusts are difficult to penetrate. Therefore the mini- 
mum requirements of api conditions for use of kieppi 
by Black Grouse (7érrao fetrix) in Finland are: 

¢ When a crust which is difficult to penetrate is covered 

by less than 10 cm soft api, then Black Grouse (Zé/7ao 
tetrix) in Finland roost in trees and on the api surface, 
but not in kieppi. The total thickness of the api is not 
important. 

¢ When there is at least 4 cm of soft api over a readily 

penetrable crust (provided that the overall thickness is 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


at least 10 cm) then Black Grouse begin to roost in 
kieppi as well as in trees and on the api surface. 
¢ They roost only in kieppi when the readily-penetrable 
crust is covered by a layer of soft snow at least 10 cm 
thick and the total thickness of the api is about 27 cm. 

¢ Marjakangas (1986) concluded that Black Grouse roost 
in open burrows on the api surface as well as in trees 
when the total thickness of the api is from 10 to 26 cm 
and exclusively in kieppi when the thickness is 27 cm 
or more. 

In his exhaustive study of the winter ecology of 
Capercaillie (7etrao parviventris) and Black Grouse in 
Finland, Seiskari (1962) put the use of kieppi into 
context of habitat type, food supply and weather. The 
temperature within Willow Grouse (Lagopus lagopus) 
kieppi rose to at least -6° even though ambient tempera- 
ture fell to -35°; relative humidity was always saturated 
and CO, density was never elevated above ambient 
(Korhonen 1980a). Andreev and Krechmar (1976) 
found kieppi temperatures of -10° and -11° with ambi- 
ent temperatures of -45°. Volkov (1968) had found simi- 
lar conditions with Hazel Grouse (Bonasia bonasia) and 
Black Grouse (7éf7ao fetrix). 

Other, smaller, birds may use kieppi also: Yellow- 
hammer (E7beriza citrinella) (Jarvi and Marjakangas 
1985), Common Redpoll (Carduelis flammea) and 
Greyheaded Tit (Poeci/e cinctus) (Korhonen 1980b), 
Snow Bunting (Plectrophanax nivalis) (Thiede 1982), 
Song Sparrow (e/ospiza melodia) (McNicholl 1980), 
Marsh Tits (Poecile palustris),Willow Tits (PR mon- 
tanus), Great Tits (P major), Long-tailed Tits (Aegsr- 
halus caudata), Snow Bunting (Plectrophanax nivalis), 
Bullfinch (Pyrrhula pyrriutla), Common Redpoll (Car- 
duelis flammea) (Novikov 1972) and others (Sulkava 
1969; 1989). Cade (1953) reported Carduelis flammea 
burrowing into api not only for thermal protection but 
to find seeds under the surface of the api. 

In periods of extreme low temperatures some birds 
such as the Redpolls (Cavduelis flammea and C. horne- 
anni) continue to feed in the coldest air zone, (on the 
api surface), by contracting their feet and legs within the 
feather coat and, using their wing primaries and tail 
feathers, support and “roll” themselves over the api sur- 
face (Johnson 1954). 

Instruments and techniques associated with thickness 
will be found later under the section entitled “General 
Procedures.” 


Qali 

Taiga snow occurs in two phases: api [ah-PEE] or 
snow on the ground and qali [KAH-lee] or the snow on 
the trees (Pruitt 1958). In temperate regions qali is of 
only transitory aesthetic importance or when determin- 
ing total water content. In the taiga, however, qali is a 
long-lasting and significant ecological factor. It has been 
an influence in the evolution of the shape of spruce 
trees; it is a powerful influence over vegetation type 
because it governs some aspects of forest succession 
(Gill et al 1973; Pruitt 1958) and it can affect human- 


made structures such as powerlines by breaking them. — 


2005 


PRUITT: WHY AND How To STUDY A SNOWCOVER 121 


FiGuRE |. Closeup photo of subnivean space and pukak crystals. Ruler ticks are 1 mm apart. Manitoba, Agassiz Provincial Forest, 4 
March 1978. Photo by Wolf Heck. 


Qali can be quantified by means of “‘qalimeters,” but the 
study of qali is in its infancy at present and the stan- 
dardization of observations would be premature (Pruitt 
1973). Research continues at Taiga Biological Station 
on characteristics and suitability of different types of 
qalimeters. 

Qali forms best under conditions of frequent, light 
snowfalls, reduced incoming solar radiation and no 
wind. A superficially-confusing phenomenon called 
kanik [KAH-nik] forms when relatively warm, mois- 
ture-laden air strikes cold objects (Pruitt 1984a, Figure 
5). It forms best under conditions near freezing, with 
light winds. Qali forms in varying amounts on horizon- 
tal surfaces, while kanik forms a layer of more-or-less 
uniform thickness, or sometimes spikes or needles, 
usually on vertical surfaces (Miller 1962, 1964, 1966). 
To complicate things, kanik sometimes forms on previ- 
ously-deposited qali or even on the api or upsik surface. 
Kanik that forms on vertical surfaces is known in 
Finnish as huurre [HOO-rre]; on horizontal surfaces it 
is known as kurra [KU-rra[ (Sirpa Rasmus, personal 
communication). 

Qali affects animals such as birds and arboreal mam- 
mals by interfering with their feeding and travel. During 
periods of heavy qali accumulation birds such as Pine 
Grosbeaks (Pinicola enuclearor), Chickadees (Poecile 
spp.) and Red Crossbills (Loaxwa curvirostra) forage on 
windy hilltops, where qali is blown off the trees. Pine 


Marten (Martes martes) and the Red Squirrel (Sczurus 
vulgaris) in Eurasia (Pulliainen 1973) and the North 
American Red Squirrel (7amzasciurus hudsonicus) find 
their arboreal activities affected by heavy gali accumu- 
lation (Pruitt, unpublished data). On the other hand, 
some small birds such as Tits and Chickadees protect 
themselves from excessive radiant heat loss by huddling 
under lumps of qali (Steen 1958). Showshoe Hares 
(Lepus americanus) use snow caves formed under qali- 
bent shrubs; thus their body radiant heat will not be lost 
to the infinite heat sink of the night sky (Pruitt 1984a). 
In contrast, qali bends shrubs over, so that their tender 
growing tips are brought within reach of the hares. This 
presents the hares with a supplementary source of 
food (Bider 1961). 


Hardness 

The snowcover that accumulates under windless con- 
ditions will be made up of flakes supported by their arms 
and touching each other only on the tips. Consequently 
most of the mass is air with relatively little ice in it. In 
the taiga or northern coniferous forest the snow season 
is characterized by little wind, a marked reduction of in- 
coming solar energy and few incursions of mild mar- 
itime or tropical air masses. The result is a snowcover 
that arrives early in the autumn and lasts all winter, rela- 
tively unaffected by thaws or wind. 


122 


Hardness (H) is an expression of the force required 
to collapse the structure of the api. Most measurements 
of this force have been presented in detailed snow ecol- 
ogy publications as gm cm”. In the remainder of this 
paper such units will be followed by S\svéme /niterna- 
tionale (S1) units such as Kg m? or Newtons [N; 5Kg = 
SON] set off by brackets. 

Hardness of taiga api can be from ~<2 to ~50 gm 
cm?[ 0.02 to 0.5N]. If the flakes are windblown they 
will be jumbled about, their fragile arms broken off and 
stripped and, when they come to rest, they will lie snug- 
ly against each other. Here they will also undergo meta- 
morphosis. A snowcover of jumbled, wind-tossed and 
broken flakes will contain more ice. The density will 
still be only about 0.30 [300 kg m=], however. Such 
a snowcover is properly termed upsik [OOP-sik]. 
Hardness of upsik can get as high as 99 000 gm cm”. 
[990 N]. Hardness is of considerable importance to 
supranivean animals as large as Caribou because it can 
impede their movement as well as their access to sub- 
nivean vegetation (Fancy and White 1985) except under 
very special conditions (Pruitt 1979) 

The upper part of the snowcover in a tundra or cold 
steppe region consists of two phases: the consolidated 
mass (upsik) and above it the moving snow, called 
siqog [see-KOK], propelled by the wind. Sigoq periodi- 
cally becomes consolidated into a sequence of drifttypes 
(Pruitt 1966, 1984a, 1999). The several types form and 
are eroded away as they reform and move over the sur- 
face of the upsik. The drift succession is known reason- 
ably well (Pruitt 1970) but any effects of the drifts on 
animals (especially subnivean animals) are poorly 
understood. The best-known effect is that of anyemanya 
[ahn-ye-MAN-ya] around obstructions to the flow of air 
(Pruitt 1984a). Sulkava (1964) has shown that anye- 
manya are important in the ecology of Grey Partridge 
(Perdix perdix) and European Hare (Lepus europaeus) 
on the Ostrobothnian Plain of western Finland. Pulli- 
ainen and fivanainen (1981) showed how distribution 
of snowcover, especially anyemanya, affected the win- 
ter diets and grit-gathering of Willow Grouse (Lagopus 
/agopus) in far northern Finland. Anyemanya are well- 
known in the folk knowledge of northern peoples as 
places where animals congregate on the exposed vege- 
tation or soil to obtain food or grit for gizzards. Such 
places are good sites for traps or snares. I have followed 
Arctic Fox (A/opex /agopus) as they zig-zagged across 
the tundra landscape, investigating one boulder-centred 
anyemanya after another; there is always the possibility of 
surprising a ptarmigan gravelling there. 

Upsik occurs in two facies which have biological 
importance (Pruitt 1984b). Convex ground surfaces, 
blown clear of snow winter after winter, are called 
vyduvi [vih-DOO-vih] and are subject to extreme cry- 
opedological processes. Concave ground surfaces col- 
lect snow each winter, are called zaboi [ZAH-boy] 
(Table 1), and are protected from temperature extremes. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Zaboi can be regulators of mesic habitats in an 
expanse of otherwise rather xeric conditions. Zaboi col- 
lect siqoq from over a wide area and concentrate it (as 
well as any contaminants attached to the siqoq particles). 
As the zaboi slowly melt during the summer, the melt- 
waters and the contained contaminants, such as radioac- 
tive particles from fallout, collect downhill of the zaboi 
(Osburn 1963). The meltwater nourishes wet sedge mats 
which absorb large quantities of it. Any contaminants, 
radioactive or otherwise, are immobilized by the sedges. 
The sedge mats are not perfect sinks, however. Eventual- 
ly the sinks will “fill up” and the contaminants will flow 
through and enter the meltwater system. This situation 
deserves attention and further research because virtually 
every city in western and central North America gets its 
water supply, originally, from zaboi (Osburn 1963). 

Small taiga mammals such as the voles Cle“iriono- 
mys spp.or Microtus spp.or shrews Sorex spp., when 
unable to escape from the api surface during cold peri- 
ods, can survive such conditions only a few hours. 
Undisturbed taiga api can have a vertical hardness range 
of <3 to 50 gm cm? [0.02 to 0.5N]. A trail made by a 
dozen passes of humans on skis can have a vertical 
hardness range of 25 to 7 500 gm cm” [0.25 to 75N], 
but a trail made by only two passes with one person on 
a snowmobile can have vertical hardnesses of 550 to 
7 000 gm cm? [5.5 to: 70N]. Moreover, a snowmobile, 
because of its weight and the churning action of the 
track, destroys the pukak layer and makes the api the 
same hardness throughout its total thickness (Pruitt, 
unpublished data). In effect, snowmobiles change api 
into upsik. Small mammals, when tunnelling through 
the layers of this highly modifed snow cover, can tum- 
ble into such trails and be unable to excavate a re-entry 
hole into the api. 

It is clear from this discussion that it is not only the 
thickness or hardness of the api that can be critical but 
also the location of the hard layer in the snowcover. 


Hardness Measurements 

I have found that a kit of snow instruments should 
have three hardness gauges: a high-range one, with a 
red stripe around it (calibrated from O to 100), a 
medium-range one (calibrated from 0 to 10) and a very- 
low-range one (calibrated from 0 to 2). There should be, 
as well, four separate discs. The discs are snapped onto 
the push-rod with their smooth faces outward. The 
largest disc results in the reading on the calibrated end 
of the push-rod to be multiplied by one, the next smaller 
disc reading is to be multiplied by 10, the next smaller 
disc is to be multiplied by 100. The smallest disc is the 
cylindrical end of the push-rod itself and its reading 
should be multiplied by 1 000. In the snow kit there 
should also be an elongated narrow bar or strip that has 
the same effective area as the largest disc. Use of this 
bar enables one to sample relatively thin layers in the 
api, thus making the results more precise. One can dif- 


2005 


ferentiate hardnesses of, for example, six layers in a 
given profile instead of only four layers with the cir- 
cular discs alone. 

Start measuring hardness by using the medium range 
gauge and attaching to it the largest disc that will fit the 
topmost layer of the api exposed on the vertical cut face. 
Push the gauge and disc, horizontally, slowly and 
steadily against the cut face. Be sure to engage the disc 
and the cut face with a horizontal motion otherwise 
error will be produced (e.g. Brown and Theberge 1990). 
At first, there will be a bit of surface crumbling. Con- 
tinue pushing until the crystalline structure suddenly 
collapses. The ratchet on the push-rod keeps the calibra- 
tions in view so that the maximum force needed can be 
read and recorded. 

Repeat the procedure several times, each in an adja- 
cent, fresh spot of the layer. Continue the procedure for 
each layer of the api. Undoubtedly you will have to use 
several combinations of gauges and discs to get combi- 
nations that will fit each layer and its hardness. Consid- 
erable practice is necessary in order to achieve consis- 
tently-reproducible results. 

A characteristic of the snowcover which I have found 
useful when considering supranivean animals is vertical 
hardness (VH). Use the proper combination of gauges 
and discs and bring them vertically down onto the sur- 
face until the crystal structure of the surface layer col- 
lapses. Horizontally cannot be substituted for vertically 
in this measurement (Pruitt 1990). A useful variant of 
VH is vertical hardness to track depth (VHT). Choose 
a disc approximating the animal track in area and push 
it vertically down to the same depth below the surface 
of the snowcover that the track reaches. The Swiss 
“ramsonde,” because of its total mass, cannot be used 
on soft taiga api. 


Density 

Density is a function of the complex history of a 
snowcover, from the aerial formation of the original 
crystals, the amount of fracturing and any possible 
thawing events they have undergone. The more com- 
plex the history, the denser the snowcover probably is. 

Density is relatively easy to measure, even using 

simple “kitchen hardware.” The weight of a given vol- 
ume of snow is divided by that volume. The expression 
(D) is a ratio, expressed without units, of the amount of 
ice in relation to the total enclosed volume. Density of 
the api has three main effects on living things: 

* It expresses the water content of the snowcover, which 
can influence the magnitude (and therefore the quality) 
of spring runoff. Increases in density, for example, 
dramatically increase the water content. 

* Density can be important to the survival of subnivean 
organisms because it governs the insulative value of the 
snowcover. Increases in density dramatically decrease 
the insulative properties of the snowcover. Density of 
taiga api runs from about 0.05 (50 kg m°) to 0.15 
(150 kg m*). It is very light and fluffy. In contrast, densi- 
ty of tundra upsik can run from about 0.15 (150 kg m*) 
to 0.5 (500 kg m°). 


PRUITT: WHY AND How To STUDY A SNOWCOVER 


VSI=1,775.5 

LogVSI=3.249 

no caribou sign 

D H VH VHT 
1 
50 
needles and debris 
air = -20 


mosses and lichens 


FIGURE 2. Sample data form and typical data from an api sta- 
tion. Station 80. 1958-02-26 Saskatchewan. 56°34'N 
108°16'W. 10m from North Shore, Lake. Mature white 
spruce-birch. Qali breaks. 


¢ The insulating properties of taiga api result in the pukak 
layer being markedly warmer than the upper layers. This 
warmth allows animal and bacterial activity to proceed. 

Such activity releases carbon dioxide in the pukak space. 

If the api contains dense layers the upward flow of air 

can be retarded and the CO, can accumulate in concen- 
trations up to 5 times ambient. This can result in behav- 
ioural changes in small mammals (Penny 1978; Penny 

and Pruitt 1984). These behavioural changes some- 
times result in Clethrionomys gapperi shifting their home 
ranges away from the affected site and returning later 
when the subnivean CO, falls to ambient concentrations. 

For determining density, choose an undisturbed verti- 
cal face of the api, brush and delimit the layers, measure 
and record thicknesses. Start with the topmost layer. 
Take the Swedish density box (preferred design) by the 
handle, hold the box horizontally and flip the end cover 
up. Push the open end of the box firmly into the api hor- 
izontally until the box is full. Be careful not to overfill 
or force more api into the box exceeding its measured 


124 


volume. Flip the end cover back over the end of the 
box. Pull the box out, scrape off any clinging snow, 
hang the box by the handle from the scales and note the 
weight. Be sure you suspend the scales by the top ring, 
not by holding the barrel. Subtract the tare of the box 
to get the weight of the enclosed snow. Divide this by 
1000 to get the density. You may want to add some coils 
of wire solder to the handle in order to bring the tare to 
a figure easily divisible by 1000. 

The Swedish density box is usable for densities up 
to about 0.25 (250 kg m® ). For determinations above 
this density there are available strong, polished metal 
tubes, sharpened or cut into saw teeth at one end and 
with two small holes so that a wire loop can be attached 
for hanging the tube from the scales. Do not use a ham- 
mer or sledge to drive a tube into the cut face of a very 
hard profile because continued use will burr the butt 
end; carry a block of wood to use as a bumper or shield 
between the end of the tube and the driver. 


Other Data to Record 

Other characteristics useful to record are: temper- 
ature of the air, temperature of each layer, temperature 
of the pukak, grain type and size for each layer, sub- 
nivean plants and, of course, the standard records of 
date, time, weather, exact locale (UTM and/or GPS 
loci), vegetation type, substrate, under tree canopy or 
not, animal activity, name of observer. 

I have found that the sample data form in Figure 2 is 
easy to use in the field. Make a master sheet with four 
replicates of the form, photocopy it as many times as 
required. One may use the 4x sheets held in a clip board 
or cut the individual forms apart with a paper cutter 
and keep them, in order, in a spring-loaded notebook. 

One does not need to establish full api profiles. 
Some aspects of animal and plant winter ecology can 
be made clear with well-chosen hardness readings. 
For example, I observed (Pruitt 1984b) how Col- 
lared Lemmings (Dicrostonyx groenlandicus) made 
their tunnels through hard tundra upsik by excavat- 
ing along a relatively soft layer floored and roofed 
with harder layers. It is clear that detailed knowl- 
edge of the natural history of the subject species will 
enable one to focus on critical features influencing 
its winter survival. 

A “Snow Index” incorporates natural history and 
behavioural characteristics and correlates them with 
movement and behaviour of the species being consid- 
ered in relation to several characteristics of the snow 
cover. For example, Pruitt (1959, 1981) showed that the 
distribution of 2. 4 gvoenlandicus on their winter range 
in northern Saskatchewan and south-central North- 
west Territories could be correlated with and predicted 
by certain characteristics of the snow cover. Henshaw 
(1968) confirmed Pruitt’s thresholds of sensitivity to 
the thickness and hardness of the snow cover in #. £ 
gran in northwestern Alaska. Stardom (1975) showed 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


how the winter activity of 2. £4 caribou in southeastern 
Manitoba correlated closely with nival factors. Stardom 
also showed that the threshold of sensitivity to thickness 
was greater in 2. 4 caribou than in RF. 4 groenlandicus. 
Darby and Pruitt (1984) put the winter movements of 7. 
t cartbou in the southeastern Manitoba taiga into per- 
spective for that mid-continent taiga region. 

Pruitt (1959) noted that the ideal nival winter range 
for XR. 4 groenlandicus had api that was thin, of little 
hardness and density, and that had not been affected 
by invasions of warm, moist air. In other words, their 
optimum winter habitat was in a cold, continental cli- 
mate that enabled the heat and moisture to flow un- 
interruptedly from the earth through the api to the 
cold, dry air above. Deviations from this ideal situa- 
tion result in worsening of over-wintering conditions 
for Caribou. 

I combined hardness and thickness data (Pruitt 1979, 
1981, 1989, 1990, 1992) of several layers in different 
combinations (based on several known aspects of Cari- 
bou winter ecology and behaviour) to arrive at what I 
call the Varrid Snow Index (VSI): 

VSI = (H>H H b Tb + Vs Ts + HhTh ) T ta/ 1000 

Where H>H = Hardness of hardest layer more than half- 

way between the substrate and the top of the snowcover. 

Hb Tb = Hardness times thickness of basal layer. 

VsTs = Vertical hardness of surface layer times thickness 

of surface layer. 

Hh Th= Hardness times thickness of hardest layer (if not 

H. iP). 

(If basal layer is the hardest, then term Hh Th drops out.) 

Tta = Total thickness of the api. 

Most records of api hardness in relation to Caribou 
have been recorded in gm cm. Pennycuick (1974) pre- 
sented factors to convert to SI units. 

This index models the relations with api of several 
subspecies of Rangifer tarandus. (R. t. tarandus |Pruitt 
1979], R. 4 groentlandicus {Pruitt 1981], 2. 4 fennicus 
[Pruitt 1989]; 2. 4 caribou (Schaefer 1990]) in Scandi- 
navia and North America. Schaefer (1996) and Schaefer 
and Pruitt (1991) used these techniques to determine 
different qualities of Woodland Caribou winter range 
in Manitoba. 

Raine (1983, 1987) devised a Snow Index that mod- 
elled the relations of Marten (artes americana) and 
Fisher to the surface and upper layers of the api in 
the taiga of southeastern Manitoba. He found that these 
two closely-related sympatric species utilized the habi- 
tat differently, governed in large degree by the charac- 
teristics of the api. Fisher movements were restricted in 
midwinter by the presence of thick, soft api. At this 
time of the year Fisher travelled on trails made by 
Snowshoe Hares (Lepus americanus) and also their 
own trails more than at other times during the snow sea- 
son. In midwinter, Fisher tended to walk through api 
and to leave a body drag. On the other hand, Marten 
appeared to be unhindered by soft api to the degree 
Fisher were. Marten tracks never showed a body drag 


2005 


PRUITT: WHY AND How To STUDY A SNOWCOVER 125 


FiGuRE 3. Technique of establishing a standardized survey of api thickness. Xs signify sites of actual thickness measurements. 


in the api. They did use Hare trails and their own trails 
in midwinter but not to the extent Fisher did. 

Raine’s Snow Index (SJ) for the top 10 cm of the api, 
in relation to Marten and Fisher is: 
ei= 1000-2 (Tx H) 

Where T = Thickness of the api in centimetres 
H = Hardness of the top 10 cm in gm cm” 

Both species were usually supported by H of 100 gm 
cm?[1 N]. This value was taken as the maximum hard- 
ness of any layer; therefore 1000 was the maximum 
value of the sum of the products. Any SI of near zero 
meant that both species could travel freely on the api 
surface, whereas an SI of 900 or more would perhaps 
indicate a hindering effect on their movements. Raine 
found that this SI modelled the responses of both 
Marten and Fisher to changes in api thickness and hard- 
ness. He also found that Fisher responded to increased 
SI by changing their gait while Marten were much less 
liable to. 

There are two notable points brought out by Raine’s 
study: 

¢ Although these closely-related species used similar 

habitats in the same region they exhibited differences 
in habitat use associated with different responses to the 
api, thus allowing the two species to co-exist 

¢ Their different responses to characteristics of the api 

enabled them to parcel out the food resources. Marten, 
being smaller and with less mass, are more subnivean, 
more arboreal and are more efficient predators on small 
mammals. Fisher, on the other hand, are better adapted 
to hunting larger prey such as Snowshoe Hares, Porcu- 
pines (Everhizon dorsatum) and ground-dwelling birds; 
indeed, they occasionally prey on Marten 

Raine’s study also demonstrated how detailed exami- 
nation of the snowcover can clarify fundamental biolog- 
ical relationships. For example, a more parsimonious 
explanation of the mid-winter differences in habitat and 
behaviour of these two closely-related species is as 
| energy-saving reactions (Formozov 1946) rather than 
curiosity, territorial or sexual interest as postulated by 
Marshall (1951) or Pulliainen (1980). 


General Procedure 

Sites for api profiles must be chosen according to cri- 
teria generated by the experimental design. For exam- 
ple, if the study is to test a null hypothesis “Snowcover 
(api) characteristics have no relation to Caribou over- 
wintering sites” then one must sample the study 
region/area with two types of snowcover sites: “con- 
trols” which are spaced over the entire region either on 
a systematic grid (by relative area of available vege- 
tation types) or by a randomization scheme, and “exper- 
imentals” which are related to actual Caribou feeding 
craters, resting sites or movement trails. The number of 
each is determined by the variance of the readings and 
the level of accuracy desired. It is as important to know 
the conditions where Caribou are o/ as to know where 
they are. 

Thickness can vary widely. The more observation 
points on your study or sampling sites usually the better. 
One way of determining thickness is to establish perma- 
nent transects with permanent markers. For example, on 
a transect establish (before snow flies) “quadrats” of 
nine sampling points, points 5 metres apart in three 
rows of three points each (Figure 3). Mark each point 
with a thin wand having alternating bands of contrasting 
colour each 10 cm high plus a bottom section long 
enough to anchor the wand firmly into the substrate. 
Paint or mark the decimeter labels, oriented so that all 
can be seen from one spot outside the quadrat. Establish 
another quadrat nearby; this will be used for destructive 
sampling (api pits) for hardness, density and crystal 
characteristics. It should also be marked so that one is 
not liable to re-use the same pit for more than one sam- 
pling. Repeat such 9-point sampling sites every 50 m 
along the permanent transect (Figure 3). 

Approach the thickness and hardness/density sites 
from the same direction each visit; this ensures that you 
do not attempt to resample the same site again. A 
wand or two will help to keep the sites differentiated. 
Frequently take record photos of the site. As a general 
rule, one can never take too many record photos. Use a 
standardized identification scheme to ensure the photos 
will be recorded correctly. 


126 


Mark your ski poles with rings of brightly coloured 
reflecting tape each 10 cm apart beginning at the tip of 
the hand grip. Use a pole, thrust handle-down, through 
the api to measure its thickness every 5 or 10 “ski steps” 
periodically as you move between sites. 

If your study region is windblown (e.g., tundra or 
steppe) determining thickness may be difficult. Very 
frequently it can be impossible to thrust a ski pole, han- 
dle first, into the hard snowcover. You may need to carry 
a thin, sharpened metal rod marked in decimeters to 
penetrate the wind-hardened snowcover. In some 
instances you may need to dig a series of small pits to 
ground level in order to expose profiles of the snow- 
cover. 

An api profile site is usually excavated so that the 
vertical cut face of the pit in the snowcover is down- 
wind and oriented so that the sun shines on it at about 
a 45° angle. This latter orientation is to emphasize the 
shadow relief of the snow crystals for photographic 
purposes. Differentiate and emphasize the layers of the 
api by gently stroking up and down with a wide, soft 
brush. Measure and record the thickness (T)(cm) of 
each layer of the api. 


a When Using a Snow Kit: 
Be sure to return instruments and discs to their proper 
place immediately after use. If you drop one into the api 
you may not find it until spring. 

¢ Record your data immediately; don’t trust your memory. 

¢ Keep the kit cold so that the instruments are at ambient 

temperature when you use them. 

¢ If snow gets inside the kit, brush out as much as you 

can. Carry a small hand-operated sucker pump in order 
to clean out all the snow. Take the kit into a warm, dry 
place, open it up, disassemble it and give it a chance to 
thaw and dry overnight. 

¢ With ambient temperatures between +5° and -5° snow 

may stick to metal parts and later freeze them immo- 
bile. Dry off the push-rod, scales, density box, etc. fre- 
quently under these conditions. 

¢ Avoid touching the metal parts with bare hands. In 

warm weather your hands heat up the parts while in 
cold weather you can get frost-nipped from handling 
the cold metal. 

The Snow Kit has had a long, evolutionary history 
from its original form (Klein, Pearce and Gold 1950). 
It continues to undergo changes. I welcome comments 
and suggestions as to how it can be modified to be even 
more useful. The actual instruments and tools needed 
will vary with the proposed research. For instance, I 
have a “complete” kit of instruments and tools for gen- 
eral research and teaching. I have found that density is 
of little or no consequence to Aangifer movements but 
hardness definitely is. Therefore, in my “caribou kit” I 
have only thickness and hardness instruments and those 
for temperature and crystal size as well as type refer- 
ence photos. 

One source of these materials is from the Science 
Shop at Lakehead University, 955 Oliver Road, Thun- 
der Bay, Ontario, Canada P7B 5E1 or “Ed Drotar” 
<ed.drotar@ lakeheadu.ca> 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


All the instruments and tools will fit into various 
slots, holes and grooves in a block of fine-grain styro- 
foam inside a bright orange angler’s “dry box,” the lid 
of which is snubbed shut with a short length of bungee 
cord. My box fits nicely inside my rucksack. A short- 
handled, flat-bladed aluminum shovel is lashed to the 
ice-axe loop of my rucksack. With a hunk of cheese and 
black bread, a thermos of tea and my Lappish bush skis 
(as well as field notebook and camera) I am outfitted for 
a day of snow study. 

So we see that using words from cultures more close- 
ly associated with snow than our relatively poverty- 
stricken English opens a different world to our view. 
Not only does specialized, precise terminology open a 
new world for us, but specialized measurements of 
characteristics of the snow cover such as duration, 
thickness, hardness and density add precision to our 
observations. These specializations mean greater under- 
standing and appreciation of adaptations of mammals 
and birds (as well as plants) to a phenomenon (snow) 
characteristic of a significant portion (winter) of the 
annual cycle of the seasons. 

It is also clear that the study of snow ecology will 
enable one to make significant contributions to the natu- 
ral history of all animals and plants of northern regions. 
The tools are available, only the thematic shift 1s 
required. 


Acknowledgments 

About sixty classes of undergraduate and graduate 
students have participated in (suffered through?) long 
days of repeating snow stations at Taiga Biological 
Station during winter field trips in Mammalogy and 
in Boreal Ecology. Many of these students have gone 
on to their own research and teaching and have con- 
tributed to our knowledge of winter ecology of mam- 
mals and birds of the taiga and tundra. I am grateful to 
Spencer Sealy and James Hare for critical comments 
on an early draft of this paper as well as to two anony- 
mous reviewers of the penultimate draft. Spencer 
Sealy also guided me through recent changes in the 
scientific names of some northern birds. My daugh- 
ter, Cheryl Ann Pruitt, has contributed her excep- 
tional editorial skills to clarify aspects of this report. 
As always, my wife and field companion, Erna, has 
kept the whole operation from falling apart. Most 
“Official” organizations and funding agencies have 
shown a remarkable lack of interest in supporting 
detailed winter ecological field studies in the taiga 
and tundra but Canadian Wildlife Service, Varrid Sub- 
arctic Research Station, Oulanka Biological Station 
and the Taiga Biological Station Research Trust have 
supported parts of this research, for which I am deep- 
ly grateful. 

Visit the Taiga Biological Station Home Page (www. 
wilds.mb.ca/taiga) for references to other reports on 
the effects of snowcover on taiga animals and plants. 


2005 


One of the outside reviewers of this paper has made 
a significant suggestion, because he has become 
“increasingly dissatisfied with phrases such as “win- 
ter ecology”, “snow ecology...” which present syntax 
problems such as “...Ecology of plants and animals in 
winter/snow...” He suggests a new word-combination 
to cover “the study of snow” on a par with “limnology”, 
“entomology” and so on. This word-combination would 
be “chionology” and made up of “chion” (Greek for 
“snow’’) and “logy” ( Greek for “knowledge of.”) This 
word-combination would fit in nicely with the three 
words “chionophile’”, “chioneuphore”’, and “chiono- 
phobe” introduced by the great Russian naturalist (and 
“chionologist’”) Alexander Nikolaevich Formozov. I 


agree with the suggestion. 


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THE CANADIAN FIELD-NATURALIST 


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Received 8 December 2003 
Accepted 7 March 2005 


Notes 


Découverte de la Salamandre a quatre orteils, Hemmidactylium scutatum, 
a Québec, Québec : limite nord-est de l’espéce sur la rive nord du 


fleuve Saint-Laurent 


DANIEL Pou.iot! et JEAN-FRANCOIS DESROCHES? 


‘Laboratoire d’écologie des eaux douces, Université du Québec a Trois-Riviéres, 3351 Boulevard des Forges, C.P. 500, 


Trois-Riviéres, Québec GOA 5H7 Canada 


*Collége de Sherbrooke, Département des Techniques d’ écologie appliquée, 475 du Parc, Sherbrooke, Québec JIK 4K1 Canada 


Pouliot, Daniel, et Jean-Francois Desroches. 2005. Découverte de la Salamandre a quatre orteils, Hemidactylium scutatum, a 
Québec, Québec : limite nord-est de l’espéce sur la rive nord du fleuve Saint-Laurent. Canadian Field-Naturalist 119(1) : 


129-131. 


En 1999 trois spécimens de Salamandre a quatre orteils ont été observés 4 Québec, sur la rive nord du fleuve Saint-Laurent. Des 
recherches effectuées au méme site en 2001 ont permis de découvrir d’ autres spécimens de méme que des pontes. Ces mentions 
élargissent la distribution connue de |’espéce de 90 km vers le nord-est-est et 80 km vers le nord-nord-ouest. II s’agit vraisem- 
blablement de la limite nord-est de cette salamandre, sur la rive nord du Saint-Laurent. 


shore of St. Lawrence River. 


Le 16 avril 1999, un adulte de la Salamandre a qua- 
tre orteils (Hemidactylium scutatum) a été découvert 
dans un boisé de la ville de Québec, dans |’ arron- 
dissement Sainte-Foy (46°45'31"N; 71°19'13"O) par 
lun des auteurs (DP). Les 22 et 24 avril de la méme 
année, deux autres individus adultes ont été décou- 
verts dans le méme boisé urbain. La premiere sala- 
mandre observée était dissimulée a l’intérieur d’un 
tronc d’arbre pourri, alors que les deux autres étaient 
cachées sous des biches, directement sur le sol. Aucun 
spécimen n’avait alors été récolté. L’habitat a cet 
endroit est une forét mixte a dominance de feuillus. 
On y retrouve l’Erable a sucre (Acer saccharum), \e 
Hétre a grandes feuilles (Fagus grandifolia), \e Chéne 
rouge (Quercus rubra), ’ Erable de Pennsylvanie (Acer 
)) pensylvanicum), \e Bouleau blanc (Beta papyrifera), 
le Sapin baumier (A4ves ba/samea), \a Pruche du Can- 
ada (7suga canadensis) et le Bouleau gris (Beta 
Populifolia). Le sol est recouvert d’une abondante liti- 
ére de feuilles mortes et les débris ligneux y sont nom- 
breux. 

Le 11 mai 2001, nous sommes retournés sur le site 
jafin notamment d’officialiser la présence de I’espéce a 
| \cet endroit par l’obtention de spécimens. Une recherche 
active effectuée entre 10h00 et 12h10 dans le boisé n’a 
pas permis d’y trouver la Salamandre a quatre orteils, 
mais 16 Salamandres cendrées (Plethodon cinereus) y 


129 


Mots-clés : Salamandre a quatre orteils, Hemidactylium scutatum, aire de distribution, extension d’aire, Québec. 


In 1999, three Four-toed Salamanders were observed at Québec, north shore of the St. Lawrence River. Field searches in 2001 
allowed the discovery of another specimen as well as egg clutches. These records extend the known range of the species 90 
km north-east-east and 80 km north-north-west. This is probably the northeastern limit of this salamander’s range on the north 


Key Words: Four-toed Salamander, Hemidactvlium scutatum, distribution, range extension, Québec. 


ont été observées. C’est finalement dans un habitat adja- 
cent, une Aulnaie a sphaigne, que cing (5) Salamandres 
a quatre orteils adultes, de méme que trois (3) masses 
d’ceufs, ont pu étre trouvées. II s’agit des premieres 
observations des ceufs de Il’espéce en milieu naturel au 
Québec. Deux spécimens adultes ont été récoltés et 
déposés au collection d’amphibiens et reptiles du 
Musée canadien de la nature (CMNAR-35712). 

Les salamandres et les masses d’ceufs étaient dis- 
simulées dans des petits ilots de Sphaigne (Sphagnum 
sp.) enchevétrés dans des racines d’arbustes, principale- 
ment des Aulnes rugueux (A/zus incana ssp. rugosa), 
et entrecoupés de petites mares formées par la fonte de 
la neige. Les plantes recensées dans cet habitat sont, 
outre les aulnes, le Chou puant (Syaplocarpus foer- 
dus), le Viorne cassinoide ( Viburnum cassinoides), \e 
Viorne a feuilles d’aulne (Viburnum alnifolium), \e 
Némopanthe mucroné (Wemopanthus mucronatus) et 
dans une moindre mesure |’Erable argenté (Acer sac- 
charinum). Ce milieu correspond a un habitat de ponte 
typique pour l’espéce si l’on se fie a la documentation 
existante (Bishop 1941; Petranka 1998). Le boisé 
adjacent est un habitat utilisé en dehors de la période 
de ponte, possiblement pour l’alimentation et I’hi- 
bernation. Ce complément d’ habitats terrestres et de 
ponte est nécessaire a la présence de la Salamandre a 
quatre orteils (Bishop 1941; Blanchard 1923). 


130 


La Salamandre a quatre orteils est une espeéce dis- 
crete et difficile 4 repérer, qui a toujours suscité |’ intérét 
des herpétologistes. Méme aujourd’ hui, elle n’est con- 
nue que de peu d’endroits dans la province (figure 1) et 
y est considérée en péril (FAPAQ 2004*). Sa présence 
au Québec a été rapportée pour la premiére fois en 
1953, prés de Hull en Outaouais (Gorham 1955). 

Jusqu’a récemment, la limite de distribution de la 
Salamandre a4 quatre orteils au Québec semblait se 
situer a l’ouest de la longitude 72°, dans la zone 
comprise a plus de 180 jours de croissance par an 
(figure 3 dans Bonin 1999 *). Toutefois, en 2000, un 
spécimen a été découvert au lac du Castor, dans la 
région de Chaudiere-Appalaches, ce qui a étendu 
aire de distribution de cette salamandre de 160 km 
vers le nord-est et 145 km vers le sud-est (Desroches 
et Couture 2002). Cette mention fut la premiere en 
dehors de la zone de limite de croissance de 180 jours 
par an, au Québec. La nouvelle localité ou fut décou- 
verte cette espece en 1999, a Québec, est également 
située en dehors de cette zone. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


La région de la ville de Québec constitue la limite 
de distribution de plusieurs espéces d’amphibiens et 
reptiles (voir les cartes de distribution dans Cook 
1984: Bider et Matte 1991). A l’est de Québec I’alti- 
tude moyenne augmente considérablement, passant 
de 116ma411 m, et la saison de croissance passe de 
170-180 jours a 150-170 jours (Robitaille et Saucier 
1998). Le domaine de l’Erabliére a Tilleul est rem- 
placé par celui de la Sapiniére 4 Bouleau jaune. Toutes 
ces différences reliées aux habitats et a la température 
portent a croire que la Salamandre a quatre orteils ne 
peut pas se retrouver beaucoup plus a |’est, sur la rive 
nord du Saint-Laurent, que le site découvert en 1999 
a Québec. 

Au Canada, on retrouve cette salamandre en On- 
tario, au Québec, au Nouveau-Brunswick et en Nou- 
velle-Ecosse. Les mentions les plus nordiques au 
Nouveau-Brunswick et en Nouvelle-Ecosse ont été 
faites a des latitudes semblables a celle de Québec 
(figure 2). En Ontario, la mention la plus nordique se 
retrouve a environ 220 km plus au nord (Bonisteel 


FIGURE |. Sites ot a été rapportée la présence de la Salamandre a quatre orteils au Québec, Canada. Les cercles noirs représentent f 


les données historiques et |’ étoile concerne le site de Québec. Cercles noirs = 1 Secteur de Hull (Gorham 1955), 2 mont | 
Saint-Grégoire (Denman 1961), 3 [le Perrot (Denman 1965), 4 Covey Hill (Gordon 1979), 5 Marlington (Sharbel 1990), 
6 Laurentides, au nord de Montréal (Bider et Matte 1991; CMNAR-33778), 7 Secteur de Trois-Riviéres (Bider et Matte 
1991; CMNAR-33773 et 33774), 8 lac du Castor, Chaudiére-Appalaches (Desroches et Couture 2002). Etoile = Ville 


de Québec (le présent article). 


2005 


206 400 relies 
= * 


280 400 500 Kelomeires 


FiGuRE 2. Aire de distribution de la Salamandre a quatre 
orteils au Canada (modifié d’ apres : Cook 1984). Les 
numéros réferent aux mentions les plus nordiques pour 
chaque province. | : Ontario (Bonisteel 1973), 2 : 
Québec (le présent article), 3 : Nouveau-Brunswick 
(Woodley et Rosen 1988), 4: Nouvelle-Ecosse (Rich- 
mond 1952; Gilhen 1984) 


1973). Ceci porte a croire que la Salamandre a quatre 
orteils pourrait se retrouver a des latitudes plus hautes 
au Québec, surtout dans I’ ouest de la province. 


Remerciements 

Les auteurs remercient Nathalie Coté, Raphaél 
Demers, Ian-Erik Gosselin et Fannie Martin pour leur 
aide sur le terrain. Ils sont également reconnaissants 
envers Mario Darsigny, qui a réalisé la carte de local- 
isations des mentions au Québec, et Francis R. Cook 
pour les informations obtenues au sujet de la Salaman- 
dre a quatre orteils au Canada. 


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Salamander (Hemidactylium scutatum) in Canada. 
COSEWIC (Committee on the Status of Endangered 
Wildlife in Canada). 21 pages. 

FAPAQ. 2004. Société de la faune et des parcs du Québec. 
http://www.fapaq.gouv.qc.ca. 


NOTES 131 


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Nova Scotia Museum, Halifax, Nova Scotia. 

Gordon, D. M. 1979. New localities for the Northern Spring 
Salamander and the Four-toed Salamander in South- 
western Quebec. Canadian Field-Naturalist 93: 193-195. 

Gorham, S. W. 1955. Notes on the Four-toed Salamander 
in the Province of Quebec. Canadian Field-Naturalist 
69: 167. 

Petranka, J. W. 1998. Salamanders of United States and 
Canada. Smithsonian Institution Press, Washington and 
London. 587 pages. 

Richmond, N. D. 1952. An addition to the herpetofauna of 
Nova Scotia and other records of amphibians and reptiles 
on Cape Breton Island. Annals of the Carnegie Museum 
322 331-332: 

Robitaille, A., et J. P. Saucier. 1998. Paysages régionaux 
du Québec méridional. Les Publications du Québec. 213 
pages + carte. 

Sharbel, T. F. 1990. A range extension for the Four-toed 
Salamander, Hemidactvlium scutatum, 1 Southern Quebec. 
Canadian Field-Naturalist 105: 285-286. 

Woodley, S. F., et M. Rosen. 1988. First record of the Four- 
toed Salamander, Hemidactylium scutatum, in New Bruns- 
wick. Canadian Field-Naturalist 102: 712. 


Received 8 October 2003 
Accepted 21 February 2005 


132 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Use of a Bridge for Day Roosting by the Hoary Bat, Zasturus cinereus 


PauL HENpDRICKS!, JOSEPH JOHNSON”, SUSAN LENARD?, and COBURN CURRIER2 


‘Montana Natural Heritage Program, 909 Locust Street, Missoula, Montana 59802 USA 
*Montana Natural Heritage Program, 1515 East Sixth Avenue, Helena, Montana 59620-1800 USA 


Hendricks, Paul, Joseph Jobnson, Susan Lenard, and Coburn Currier. 2004. Use of a bridge for day roosting by the Hoary 
Bat, Lasiurus cinereus. Canadian Field-Naturalist 119(1): 132-133. 


The Hoary Bat (Lasiurus cinereus) is a migratory species with the widest distribution of all New World bats. It is a solitary 
species that roosts during the day and night primarily among tree foliage. During a survey of 130 highway structures (bridges 
and culverts) in south central Montana for evidence of use by bats, we discovered a female Hoary Bat with young in mid 
July 2003 using a wooden bridge as a day roost. This is the first report of Hoary Bats using a bridge as roosting habitat. 


Key Words: Hoary Bat, Zasvurus cinereus, roosting, habitat, bridge use, Montana. 


The Hoary Bat (Zas/urus cinereus) has the widest 
distribution of all New World bats (Shump and Shump 
1982). Across their range males and females tend to 
roost alone (Constantine 1966; Barbour and Davis 1969; 
van Zyll de Jong 1985; Nagorsen and Brigham 1993). 
Roosting individuals are encountered infrequently in 
deciduous and coniferous tree foliage, the preferred 
roosting habitat, and much remains to be learned about 
their roosting habits. 

During July and August 2003 we examined 130 
highway structures (bridges and culverts) in Carbon, 
Stillwater, and Yellowstone counties, south central 
Montana, for evidence of use by bats as day or night 
roosts. Twelve bridges were used as day roosts by four 
species of bats. Two bridges supported maternity col- 
onies of Little Brown Myotis (Avortis Jucifugus), and 
two harbored maternity colonies of Big Brown Bats 
(Eptesicus fuscus). We estimated these colonies includ- 
ed 15-90 adults; at the remaining eight bridges we 
found 1-8 individual bats. Bat species present among 
the 12 bridge day roosts included Big Brown Bat at ten, 
Little Brown Myotis at two, Western Small-footed Myo- 
tis (M. ci/iolabrum) at two, and Hoary Bat at one. 

On 16 July at about 11:00, we discovered a female 
Hoary Bat roosting in a bridge spanning Shane Creek 
(45°36'42.7"N, 109°16'48.7"W; 1125 m elevation) on 
Montana Highway 78, about 4.8 km S of Columbus, 
Stillwater County. She was wedged in a tapering 4 cm 
wide slot (ca. 20 cm deep) between two wooden girders 
on the underside of the bridge 5.3 m above bare ground 
(Figure 1); only a shallow narrow channel of water was 
flowing under the bridge at the time. Using an 8x binoc- 
ular and spotlight we could see at least two naked pups 
partially exposed and crawling under her folded wings. 
A maternity colony of about 15 adult Big Brown Bats 
occupied the same slot 1.5 m from the female Hoary 
Bat but where the slot was slightly narrower. 

The bridge was a two-lane wooden girder design 
(19.8 m long) with an asphalt deck surface, and pro- 
vided roosting sites similar to some railroad bridges 
(see Davis and Cockrum 1963). Vegetation adjacent 
to the bridge included a riparian strip of Plains Cot- 
tonwood (Populus deltoides) and willow (Salix sp./, 


and pasture intermixed with rural home development; 
nearby sandstone bluffs supported open stands of Pon- 
derosa Pine (Pinus ponderosa). We revisited the bridge 
the two days following discovery of the female Hoary 
Bat, as well as on 18 August, 9 and 16 September, but 
no Hoary Bat was present on any of these days, al- 
though about 15 Big Brown Bats continued to use the 
bridge as a day roost until the last date, when none 
was present. 

Records of Hoary Bats roosting in sites other than 
tree foliage are unusual, and use of man-made structures 
is rare. Appearance in caves is the most frequently 
reported alternative roost site, but Hoary Bats found 
in caves usually are dead or dying individuals (Mum- 
ford 1953; Beer 1954; Myers 1960). Other reported 
roosting sites include a squirrel nest, a woodpecker cav- 
ity, a tree hollow, a driftwood plank, a cellar door, and 
the side of a building (Bailey 1936; Neill 1952; Shump 
and Shump 1982; Nagorsen and Brigham 1993). 

Use of bridges by Hoary Bats for day or night roost- 
ing has not been reported previously in the literature. 
Ellison et al. (2003) mention collection of a Hoary Bat 
at a bridge, but this is based on a specimen (MSU5676 
at Montana State University, Bozeman) taken at 
“Shedd’s Bridge” in Gallatin County, Montana on 11 
September 1964; no data associated with the speci- 
men indicate that it was roosting in the bridge. The 
Hoary Bat was not among the 17 bat species found dur- 
ing a broad survey of 2421 highway structures (bridges 
and culverts) across the western and southern United 
States (Keeley and Tuttle 1999). Studies of smaller sam- 
ples of bridges in Arizona, northern California, and the 
Oregon Coast Range also did not report the presence 
of Hoary Bats (Davis and Cockrum 1963; Pierson et 
al. 1996; Adam and Hayes 2000). 

Lack of prior reports of Hoary Bats using bridges 
for roosting (Shump and Shump 1982; Pierson et al. 
1996; Keeley and Tuttle 1999) indicates that bridge use 
by this species is a relatively rare occurrence. The Hoary 
Bat is a solitary roosting species that switches roosts 
frequently (Lewis 1995); this could make detection of 
their use of any particular highway structure highly 
unlikely. Bridge surveys often involve only one or a few 


2005 


FIGURE |. Underside of wooden bridge spanning Shane Creek, 
Stillwater County, Montana, showing July 2003 day- 
roost locations of a female Hoary Bat (A) and a mater- 
nity colony of Big Brown Bats (B). 


visits to specific structures (Davis and Cockrum 1963; 
Keeley and Tuttle 1999; Adam and Hayes 2000; this 
study), or else monitor activity more intensively at a 
small number of structures (Davis and Cockrum 1963; 
Pierson et al. 1996; Adam and Hayes 2000); the num- 
ber of bridge visits or scope of intensive surveys may 
be inadequate to detect an extremely low frequency 
of bridge use by the Hoary Bat. Studies that focus on 
bridges known to be used as night roosts (Pierson et 
al. 1996; Adams and Hayes 2000) may overlook Hoary 
Bats if their use of bridges is more likely during the 
day, the time when we made our observation. Thus, 
low roost site fidelity and a solitary nature may make 
detecting the rare occupancy of bridges by Hoary Bats 
even more problematic if only single visits to bridges 
are made during daytime surveys, or studies are focused 
primarily or exclusively on night roost dynamics. 


Acknowledgments 

Our observations were made during a cost reim- 
bursement contract (MDT Project #8159) between the 
Montana Department of Transportation and Montana 
State Library-Natural Resource Information System/ 
Montana Natural Heritage Program. The opinions, find- 
ings, and conclusions expressed in this publication are 
those of the authors and not necessarily those of the 
Montana Department of Transportation or the Feder- 
al Highway Administration. We thank Susan Sillick 
(MDT) and Jim Hill (NRIS) for administering this 


NOTES 


133 


contract, and Larry Urban (MDT) for being receptive 
to the initial idea. Laura Ellison (US Geological Survey) 
identified the source of their bridge use record. Our 
paper benefited from the comments of two reviewers. 


Literature Cited 

Adam, M. D., and J. P. Hayes. 2000. Use of bridges as night 
roosts by bats in the Oregon Coast Range. Journal of Mam- 
malogy 81: 402-407. 

Bailey, V. 1936. The mammals and life zones of Oregon. 
North American Fauna 55. 416 pages. 

Barbour, R. W., and W. H. Davis. 1969. Bats of America. 
University Press of Kentucky, Lexington, Kentucky. 286 
pages. 

Beer, J. R. 1954. A record of the hoary bat from a cave. 
Journal of Mammalogy 35: 116. 

Constantine, D. G. 1966. Ecological observations on lasiurine 
bats in lowa. Journal of Mammalogy 47: 34-41. 

Davis, R., and E. L. Cockrum. 1963. Bridges utilized as 
day-roosts by bats. Joumal of Mammalogy 44: 428-430. 

Ellison, L. E., T. J. O’Shea, M. A. Bogan, A. L. Everette, 
and D. M. Schneider. 2003. Existing data on colonies of 
bats in the United States: summary and analysis of the 
U.S. Geological Survey’s bat population database. Pages 
127-237 im Monitoring trends in bat populations of the 
United States and territories: problems and prospects. 
U.S. Geological Survey, Biological Resources Division, 
Information and Technology Report USGS/BRD/ITR-2003- 
0003. Edited by T. J. O’Shea and M. A. Bogan. National 
Technical Information Service, Springfield, Virginia. 274 
pages. 

Keeley, B. W., and M. D. Tuttle. 1999. Bats in American 
bridges. Resource Publication 4. Bat Conservation Inter- 
national, Austin, Texas. 41 pages. 

Lewis, S. E. 1995. Roost fidelity of bats: a review. Journal 
of Mammalogy 76: 481-496. 

Mumford, R. E. 1953. Hoary bat skull in an Indiana cave. 
Journal of Mammalogy 34: 121. 

Myers, R. F. 1960. Lasiurus from Missouri caves. Journal 
of Mammalogy 41: 114-117. 

Nagorsen, D. W., and R. M. Brigham. 1993. Bats of British 
Columbia. UBC Press. Vancouver, British Columbia. 164 
pages. 

Neill. W. T. 1952. Hoary bat in a squirrel’s nest Journal of 
Mammalogy 33: 113. 

Pierson, E. D., W. E. Rainey, and R. M. Miller. 1996. Night 
roost sampling: a window on the forest bat community in 
northern California. Pages 151-163 /7 Bats and forests: 
proceedings of the Victoria Symposium. Edited by R. M. 
Barclay and R. M. Brigham. Ministry of Forests Research 
Program Working Paper 23/1996, Victoria, British Colum- 
bia. 292 pages. 

Shump, K. A., Jr., and A. U. Shump. 1982. Lasiurus cinereus. 
Mammalian Species 185: 1-5. 

van Zyll de Jong, C. G. 1985. Handbook of Canadian Mam- 
mals. 2. Bats. National Museum of Natural Sciences, Ottawa, 
Ontario. 212 pages. 


Received 25 March 2004 
Accepted 28 January 2005 


134 THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Aggressive Behaviour Exhibited by a San Joaquin Kit Fox, Vidves 


Mmacrous mutica 


HOWARD O. CLARK, JR. 


California State University, Stanislaus. Endangered Species Recovery Program. 1900 North Gateway Boulevard, Suite 101, 


Fresno, California, 93727-1622, USA. 


Current address: H.T. Harvey & Associates, 423 West Fallbrook, Suite 202, Fresno, California 93711-6138, USA. 


Clark, Howard O., Jr. 2005. Aggressive behaviour exhibited by a San Joaquin Kit Fox Viddpes macrotis mutica. Canadian Field- 


Naturalist 119(1): 134. 


While placing a hairtube-trapping device near a Kit Fox natal den, I observed a juvenile Kit Fox exhibit aggressive behaviour 
towards my presence. Although Kit Foxes usually are not aggressive, my observation demonstrated that some Kit Foxes defend 


their natal dens when alarmed. 


Key Words: San Joaquin Kit Fox, Vidpes macrotis mutica, aggressive behaviour, natal den, California. 


Kamler et al. (2000) reported an incident where a 
juvenile male Swift Fox (Via/pes velox) exhibited 
aggressive behaviors toward researchers. Like the Swift 
Fox, Kit Foxes are usually acquiescent and do not 
readily display aggression toward humans (when not 
trapped). Similar to the observations of Kamler et al. 
(2000), I witnessed the aggressive behaviour of a radio- 
collared juvenile Kit Fox as I was placing prototype 
hairtube traps (Clark et al. 2003) in the vicinity of its 
natal den. As I was placing the trap, the male pup 
aggressively approached me with a series of sharp 
barks. He circled around me within 3 m — sniffing and 
barking. He then stuck his head into the hairtube trap 
twice while I was watching him, but did not remove any 
bait. He continued to pace around me and bark for about 
10 min. This observation occurred at 20:45 on 25 May 
1999, along the California Aqueduct, near Lost Hills, 
Kern County, California, 35°36.98'N, 119°41.66'W 
(WGS84/NAD83). It was dark at the time; sunset was 
at 20:03. The natal den was being used by two adult 
(male and female) Kit Foxes and three male pups. All 
five Kit Foxes were radio-collared, with the juveniles 
being trapped and radio-collared on 3 May 1999. 

It appeared that the juvenile was protecting the natal 
den area and was curious to discover my reaction 
toward his aggressive approaches. Aggressive foxes that 
readily approach potential danger might experience a 
higher rate of mortality than those that select to run and 
take cover in dens. By November 1999, two of these 
juveniles were killed by Coyotes (Canis /atrans) and 


one had presumably dispersed (Clark 2001). Coyotes 
are a common source of mortality for Kit Foxes (Ralls 
and White 1995). My observation suggests that some 
Kit Foxes will defend their natal areas and aggressively 
approach potential threats to themselves and family 
members, as also described in Kamler et al. (2000). 


Acknowledgments 

The California Department of Water Resources 
gave permission to enter the California Aqueduct 
right-of-way. Funding of this note provided by the 
U.S. Department of the Interior, Bureau of Reclama- 
tion. J. Clark also witnessed the fox interaction. 


Literature Cited 

Clark, H. O., Jr. 2001. Endangered San Joaquin Kit Fox and 
Non-native Red Fox: Interspecific Competitive Interactions. 
M.S. thesis, California State University, Fresno. 54 pages. 

Clark, H. O., Jr., B. L. Cypher, P. A. Kelly, D. F. Williams, 
and S. D. Clifton. 2003. Use of a hair-sampling tube to 
detect the San Joaquin Kit Fox. Transactions of the Western 
Section of the Wildlife Society. 38/39: 29-30. 

Kamler, J. F., W. B. Ballard, and K. Mote. 2000. Aggressive 
behavior exhibited by a Swift Fox, V/pes ve/ox. Canadian 
Field-Naturalist 114: 506. 

Ralls, K., and P. J. White. 1995. Predation on San Joaquin Kit 
Foxes by larger canids. Journal of Mammalogy 76: 723- 
22: 


Received 29 March 2004 
Accepted 7 March 2005 


GN 


2005 NOTES 13 


An Ancient Wolf, Canus /upus, Den and Associated Human Activity 
in the Southwestern Yukon Territory 


RICHARD FARNELL!, P. GREGORY HARE?, and DANIEL R. DRUMMOND! 


'Department of Environment, Government of Yukon, Box 2703, Whitehorse, Yukon, Y1A 2C6 Canada 
"Department of Tourism and Culture, Government of Yukon, Box 2703, Whitehorse, Yukon, Y1A 2C6 Canada 


Farnell, Richard, P. Gregory Hare, and Daniel R. Drummond. 2005. An ancient wolf den and associated human activity in 
the southwestern Yukon Territory. Canadian Field-Naturalist 119(1): 135-136. 

The recovery of an ancient hunting artifact in an active Wolf den indicates that Wolf denning sites may be reused for many 

centuries. It also suggests that traditional practices of predator management by humans may have great antiquity. 


Key Words: Wolf, Canis /upus, den, archaeology, bow and arrow, Yukon. 


Yukon Wolves, Canis Lupus, typically dig dens in 
May to rear pups (Rausch 1967). They may occupy 
the same den for several consecutive years or inter- 
mittently over long periods; however, the long-term 
persistence of Wolf dens in the Yukon has not been 
explored. The recent recovery of an antler arrow point 
within a Wolf den provides evidence that Wolves may 
return to the same denning sites over many centuries. 

In May 1995, during a routine monitoring visit to a 
Wolf den in a remote area of southwestern Yukon, 
DRD [Daniel R. Drummond] entered an entrance tun- 
nel leading to the main chamber (Figure 1). Here, he 
found and extracted a barbed antler point that was em- 
bedded in the tunnel side-wall near the tunnel entrance. 
The projectile point is typical of a style of point used 
by Southern Tutchone hunters throughout the past 1200 
years (Hare et al. 2004) (Figure 2). The artifact was 


FIGURE 2. Photo of barbed antler arrow point (scale bar in 
centimeters). 


submitted to Beta Analytic Laboratories for AMS radio- 
carbon dating and returned a date of 850 + 50 B.P. 
(Beta- 162350). 

This date provides evidence that the den was in use 
by at least 1040 to 1270 AD (calibrated, 2 Sigma vari- 
ation). While there was significant amount of biotur- 
bation at the site, the excellent preservation of such a 
fragile, organic artifact indicates that the projectile 
point has remained in buried, dry conditions for most 
of that entire period. 

The southwest Yukon den (61°N 5', 137°W 54’) is 
known as the Kloo Lake den. It was monitored annu- 
ally for reproductive activity from 1992 to 1998 for a 
Wolf fertility control experiment (Spence 1998). The 
den characteristics are typical of those found in Yukon 
and Alaska (A. Baer, Yukon Department of Environ- 
ment, unpublished data; Ballard and Dau 1983). The 

, © den site is well suited for Wolves. It has a southern 

7 is “= exposure and is situated on an old elevated river terrace 

' composed of very stable frost-free lacustrine soils. It is 
: located in a mixed open stand of spruce (//cea spp.), 
Trembling Aspen (/opulus tremuloides), and willow 
(Sa/ix spp.). It overlooks a large valley composed most- 
ly of wetland traversed by the Jarvis River which flows 
into Kloo Lake 13 km downstream. There is substantial 
FiGuRE 1. Photo of D. Drummond examining entrance of — Prey available in the area as it is situated in close prox- 
Kloo Lake den tunnel. imity to calving and summer habitats of Woodland 


136 


Caribou (Rangifer tarandus caribou) and Moose (A/ces 
alces) (Hayes et al. 2003). Caribou in particular are 
known to have occurred in the area in large numbers 
for at least 8000 years (Farnell et al. 2004). 

There are several possible explanations to account 
for a barbed antler projectile point at this Wolf den. 
An ancient hunter may have lost the projectile point 
while “Wolf denning” or attempting to kill a wolf to 
use the hide for clothing many centuries ago. Alter- 
natively a Wolf or other predator or scavenger could 
have transported prey remains with the point embed- 
ded from a previously wounded animal. Likewise, a 
hunter could have wounded a Wolf that subsequently 
returned to the den site. There is also the possibility 
that a hunter may have inadvertently dropped the pro- 
jectile point at this location. 

It is possible that the Wolf den was recently con- 
structed on top of an archeological site that contained 
an antler point but this situation is highly unlikely. Two 
site inspections (2002 and 2003) of the Wolf den 
demonstrated that despite large surface exposures and 
sediment upheaval there was no other archeological 
evidence at the site (e.g., lithics, burned bone, charcoal, 
fire cracked rock). 

Knowledge of historical locations of Wolf dens and 
the cultural and the traditional practice of “Wolf den- 
ning” (e.g., culling or capturing Wolf pups at dens dur- 
ing spring) is documented in both Yukon First Nation 
oral history and ethnographic documentation (John- 
son 1994*; Allen 1993*; LeGros 1981*; Art Johns, 
Carcross/Tagish First Nation elder personal commu- 
nication to R. Farnell, Tom Smith, Kaska Dena elder, 
commentary in Northern Native Broadcastings, “The 
Come Back Trail” 1988; Percy Henry, Tr’on dek 
Hwech’in, elder testimony to Alaska Board of Game, 
Anchorage, Alaska 1998). If the recovery of an 850- 
year-old arrow point within a den is related to the prac- 
tice of “Wolf denning”, it indicates that it is a tradition- 
al practice of long standing. It is also possible that it 
resulted from an effort to obtain Wolf pelts for use in 
clothing. However this seems unlikely as Wolf pelts are 
in extremely poor condition during spring and summer. 

In view of the persistence of Wolf den reuse, it 1s 
possible that native hunters could have used predictable 
denning behavior to reduce interspecific competition 
for common prey. There is a large body of evidence 
that Wolves limit Caribou and Moose numbers (Hayes 
et al. 2003). At the same time, Moose and Caribou 
were critical to the survival of subarctic Southern 
Tutchone. Reducing Wolf numbers at dens would 
have improved the likelihood of success in hunting. 

Mech and Packard (1990) first reported evidence 
for possible use of a Wolf den spanning a period of 
700 years or more on Ellesmere Island, Northwest 
Territories based on fossil prey remains. On Ellesmere, 
Wolves cannot dig dens because of permafrost. The 
Ellesmere denning site was a cave and there are few 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


in the area making it a uniquely suitable location. In 
Yukon, den longevity must rely on substrate stability 
and suitable regional prey availability—conditions 
that must have persisted at the Kloo Lake den for 
many centuries. At our latitude Wolves have many 
options for places to den; yet a surprising proportion 
of dens are reused. It may be that wolves are attracted 
to places previously used by other Wolves and this is 
an important factor in the continued use of some dens. 


Acknowledgments 

The Yukon Department of Environment and Depart- 
ment of Tourism and Culture supported this study. 
Suggestions provided by Robert O. Stephenson, Alaska 
Department of Fish and Game, Fairbanks, and Layne 
G. Adams, U. S. Geological Survey, Alaska Science 
Center, Anchorage are greatly appreciated. 


Documents Cited (marked* in text) 

Allen, J. 1993. Traditional knowledge report: Aishihik caribou 
recovery area. Champaign and Aishihik First Nation and 
Yukon Department of Environment. Whitehorse, Yukon. 

Johnson, M. 1994. Aishihik and Kluane caribou recovery: 
A summary of Kluane First Nation traditional knowledge 
interviews and recommendations. Kluane First Nation and 
Yukon Department of Environment, Whitehorse, Yukon. 

LeGros, D. 1981 Structure Socio-Culturelle et Rapports De 
Domination Chez Les Indiens Tutchone Septentrionaux 
du Yukon au Dix-Neuviene Siecle. Unpublished Ph.D. 
thesis, University of British Columbia. 


Literature Cited 

Ballard, W. B., and J. R. Dau. 1983. Characteristics of Gray 
Wolf, Canis /upus, den and rendezvous sites in Southcen- 
tral Alaska. Canadian Field-Naturalist 97: 299-302. 

Farnell, R., P. G. Hare, S. Greer, E. Blake, V. Bowyer, 
and C. Schweger. 2004. Multidisciplinary investigations 
of alpine ice patches in Southwest Yukon, Canada: Paleo- 
environmental and paleobiological investigations. Arctic 
57: 247-259. 

Hare, P. G., S. Greer, R. Gotthardt, R Farnell, V. Bowyer, 
and C. Schweger. 2004. Multidisciplinary investigations of 
alpine ice patches in Southwest Yukon, Canada: Ethnogra- 
phic and archaeological investigations. Arctic 57: 260-272. 

Hayes, R. D., R. Farnell, R. M. P. Ward, J. Carey, M. Dehn, 
G. W. Kuzyk, A. M. Baer, C. L. Gardner, and M. 
O’Donoghue. 2003. Experimental reduction of wolves in 
Yukon: ungulate responses and management implications. 
Widlife Monographs 152. 

Mech, L. D., and J. M. Packard. 1990. Possible use of wolf, 
Canis lupus, den over several centuries. Canadian Field- 
Naturalist 104: 484-485. 

Rausch, R. A. 1967. Some aspects of population ecology of 
wolves, Alaska. American Zoologist 7: 253-265. 

Spence, C. E. 1998. Fertility control and the ecological con- 
sequences of managing northern wolf populations. M.S. 
thesis. Department of Botany. University of Toronto, 
Toronto, Ontario. 


Received 26 January 2004 
Accepted 21 February 2005 


2005 


NOTES 137 


Papillate Watermeal, Wo/ffia brasiliensis, in Eastern Ontario: An Addition 


to the Flora of Canada 


E. R. THOMSON 


P.O. Box 262, Merrickville, Ontario KOG INO Canada 


Thomson, E. R. 2005. Papillate Watermeal, Wo/ffa brasiliensis, in eastern Ontario: An addition to the flora of Canada. Canadian 


Field-Naturalist 119(1): 137-138. 


Wolffa brasiliensis is reported from eastern Ontario, at several sites within Frontenac and Lanark Counties. This is the first 


published record of W. brasiliensis in Canada. 


Key Words: Wol/fiia brasiliensis, Wolffia papulifera, Watermeal, range extension, new record, Ontario, Canada. 


Wolffia (Watermeal) is a small genus in the Lem- 
naceae (Duckweed Family) which consists of about | 1 
species and includes the smallest known flowering 
plants in the world. According to Scoggan (1978), 
Boivin (1967), and Dore (1957) only two species occur 
in Canada: Wolffa columbiana Karsten (reported by 
Scoggan and Boivin as W. arhiza Wimm.) and W. borve- 
alis (Engelm.) Landolt (referred to as W. punctrata by 
Scoggan, Boivin, Dore and other American authors). 
Both species occur, often together, in eastern Ontario 
(Dore 1957; Newmaster et al. 1998). A third species, W 
brasiliensis Weddell (= W. papulifera C. H. Thompson), 
has a range which extends from South America north 
to the eastern United States where it approaches the 
Canadian border (Landolt 2000). It has been found as 
far north as Berrien County, Michigan, and the shore of 
Lake Erie east of Cleveland, Ohio (Hess 1986; Dore 
19577): 

While conducting a botanical survey near Garter 
Lake, Bedford District, Frontenac County, Ontario, on 
28 June 2003, I came upon a small, shallow beaver 
pond with Wo/fia growing on the surface. The fronds 
of these plants were flattened on top as in W. Gorealis, 
but each bore a single, distinct papilla in the middle of 
the upper surface (Figure |). The apex of the frond was 
rounded in dorsal view, matching the description of W. 
brasiliensis, rather than that of the acutely pointed W 
borealis. 

No published record of W. Arasi/iensis in Canada has 
been found. (Reports of W. brasz/iensis from Ontario, 
including that of Kartesz (1999), are based on W 
Lorealis (Dore 1957; Morton and Venn 1990).) As well, 
in the DAO herbarium (acronym according to Holm- 
gren et al. 1990), no W. Gvasi/iensis was detected in col- 
lections of the closely related W. borealis, athough, as 
Dore notes, it is difficult to distinguish between them 
in the dry state. There is, however, a single specimen 
there of W. brasiliensis, collected on 10 September 1964 
by Isabel Bayly, which has interesting label data 
[DAO 717812]. Although the collection was made in 
the greenhouse at Carleton University, Ottawa, from 
“dense vigorous growth in aerated aquarium with 
Eichornia |sic]’, the original material was said to have 


FIGURE 1. Vegetative fronds of Wolffa brasiliensis from 
Frontenac County, Ontario, showing central papilla. 
Photograph by Stephen J. Darbyshire. 


come from an “artificial pond caused by a construction 
error, flooded 8-10 years previously”, near Christie 
Lake, in Lanark County, Ontario. It is interesting to note 
that Christie Lake is only about 20 km from the Garter 
Lake site. 

Situated in a predominantly deciduous forest, the 
Beaver pond near Garter Lake was edged with Wool- 
grass (Scirpus cyperinus) and Bristly Sedge (Carer 
comosa) and contained a variety of water plants includ- 
ing Watershield (4vasenia schreberi), Greater Bladder- 
wort (W/tricularia macrorhiza, Bur-reed (Sparganium 
americanum), and Coontail (Ceratophyllum demer- 
sum). Mixed in with the W. drasiliensis were a few 
plants of W. columbiana. Over the summer of 2003, the 
colony increased in size. Whereas on 3 July it had 


138 


covered an area about 10 m? of the pond’s surface, by 
16 September it covered an area of about 40 m?. The 
ratio of abundance of the two species appeared to 
remain fairly constant. 

Further surveys produced additional findings of W 
brasiliensis in the same general area of Frontenac and 
Lanark Counties (see Appendix). Six locations were 
checked where patches of Wo/fia were present. Wolffia 
brasiliensis was present in five of them, and in each of 
these five populations, it was the predominant species. 
The populations were found in a variety of quiet, shal- 
low-water habitats, including beaver ponds, cattail 
marshes, and an old, beaver wetland of grasses, sedges 
and holly. At all sites, W. columbiana was also present 
but in lower numbers. At both sites on Briggs Lane 
(collections 3 and 4), some W. Gorvea/is was also found 
mixed with the other two species. In collection 7, 
from 16 October, a few plants of W drasilrensis were 
in flower. The style with its dark stigma was clearly 
visible protruding from the floral cavity. On some indi- 
viduals, the single anther could also be seen, still within 
the cavity. 

Has Wolffa brasiliensis spread to this area recently, 
or has it been growing here unnoticed for years? The 
Bayly collection from 1964 seems to suggest the latter. 
The similarity of this species to W Gorealis could cause 
it to be easily overlooked. Judging from the several 
locations found, the farthest ones being about 20 km 
apart, it seems unlikely that it is a recent, isolated, 
chance introduction by waterfowl. Rather, it appears to 
be well established in the area. 


Acknowledgments 

I am indebted to Stephen Darbyshire of Agriculture 
and Agri-Food Canada, who provided encouragement 
and generous assistance in many ways, including 
preparing the herbarium specimens, photographing the 


APPENDIX 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


plants, locating reference material, and reviewing of the 
manuscript. 


Literature Cited 

Boivin, B. 1967. Enumération des plantes du Canada. VI — 
Monopsides, (2°me partie). Le Naturaliste Canadien 94: 
471-528. 

Dore, W. G. 1957. Wo/ffia in Canada. Canadian Field-Nat- 
uralist 71: 10-16. 

Hess, W. J. 1986. Wolfia papulifera Thompson (Lemnaceae), 
new to Michigan. Sida 11: 407-411. 

Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. 
Index Herbariorum. Part I: The Herbaria of the World. 8" 
edition. New York Botanical Garden, Bronx, New York. 
693 pages. 

Kartesz, J. T. 1999. A Synonymized Checklist and Atlas with 
Biological Attributes for the Vascular Flora of the United 
States, Canada, and Greenland. First edition. 7: Kartesz, 
J. T., and C. A. Meacham. Synthesis of the North American 
Flora, version 1.0. North Carolina Botanical Garden, 
Chapel Hill, North Carolina. 

Landolt, E. 2000. 204. Lemnaceae Gray. Duckweed Family. 
Pages 143-153 77 Flora of North America North of Mexico. 
Volume 22. Edited by Flora of North America Editorial 
Committee. Oxford University Press, New York, New York. 

Morton, J. K., and J. M. Venn. 1990. A Checklist of the Flora 
of Ontario, Vascular Plants. Department of Biology, Univer- 
sity of Waterloo, Waterloo, Ontario. 

Newmaster, S. G., A. Lehela, P. W. C. Uhlig, S. McMurray, 
and M. J. Oldham. 1998. Ontario Plant List. Forest 
Research Information Paper Number 123, Ontario Forest 
Research Institute, OMNR, Sault Ste. Marie, Ontario. 

Scoggan, H. J. 1978. The Flora of Canada Part 2 — Pterido- 
phyta, Gymnospermae, Monocotyledoneae. National Muse- 
um of Natural Sciences Publications in Botany, Number 7: 
93-545. National Museums of Canada, Ottawa, Ontario. 


Received 8 November 2003 
Accepted 23 February 2005 


Collections of Wolfia brasiliensis from Ontario deposited in the herbarium of Agriculture and Agri-Food Canada, Ottawa (DAO). 

1. Frontenac County, Bedford District, Garter Lake, UTM: 18T 0377173 4941383 [44°37'02"N, 76°32'53"W]; growing in a 
small, shallow beaver pond on stream flowing into Garter Lake; E. R. Thomson, 3 July 2003 (DAO 792125) 

2. Frontenac County, Bedford District, Garter Lake, UTM: 18T 0377173 4941383 [44°37'02"N, 76°32'53"W]; growing in a 
small, shallow beaver pond on stream flowing into Garter Lake; E. R. Thomson, 12 July 2003 (DAO 792123) 

3. Frontenac County, Bedford District, west of Wolfe Lake, Briggs Lane, UTM: 18T 0379606 4948662 [44°41'01"N, 
76°31'05"W]; very abundant around the edge of a cattail marsh; E. R. Thomson, 17 September 2003 (DAO 792124) 

4. Frontenac County, Bedford District, west of Wolfe Lake, Briggs Lane, UTM: 18T 0378843 4948442 [44°40'53"N, 
76°31'42"W]; thickly covering a shallow pond; mostly W. drasi/iensis with some W. columbiana and a few W. borealis, 
E. R. Thomson, 24 September 2003 (DAO 792121) 

5. Lanark County, south of Christie Lake on Althorpe Road, 200 yards west of Hanna Road, UTM: 18T 0385598 
4959533 [44°46'56"N, 76°26'43"W]; cattail marsh / pond; E. R. Thomson, 26 September 2003 (DAO 792122) 

6. Frontenac County, Bedford District, west of Wolfe Lake, UTM: 18T 0379874 4949264 [44°41'16"N, 76°30'52"W]; small 
patches of open water in an old beaver wetland of grasses, sedges and holly; E. R. Thomson, 26 September 2003 (DAO 
792120) 

7. Frontenac County, Bedford District, west of Wolfe Lake, UTM: 18T 0380368 4950143 [44°41'46"N, 76°30'33"W]; shallow 
old beaver pond; mostly sterile fronds, a few fertile fronds; E. R. Thomson, 16 October 2003 (DAO 792258) 


2005 NOTES 139 


Record Size Female Coyote, Camis latrans 


JONATHAN G. Way! and RoBERT L. PROIETTO? 


'Urban Ecology Institute and Biology Department, Boston College, Higgins Hall, Chestnut Hill, Massachusetts 02467; USA; 
e-mail: wayjo@bc.edu 
Biology Department, Boston College, Chestnut Hill, Massachusetts 02487 USA; e-mail: robertproietto@hotmail.com 


Way, Jonathan G., and Robert L. Poietto. 2004. Record size female Coyote, Canis /atrans. Canadian Field-Naturalist 119(1): 
139-140. 


On 11 March 2004 we recaptured and re-radio-collared an 8-9 yr old, 25.1 kg (55.3 Ib), 157 cm long (tip of nose to tail tip) 
female Eastern Coyote (Canis /arrans) in the town of Barnstable on Cape Cod, Massachusetts, that was originally captured in 


November 1998. This is believed to be the largest female Coyote ever recorded 


Key Words: Eastern Coyote, Canis /atrans, body size, Cape Cod, Coyote, Massachusetts, urbanization, weight 


Coyote (Canvs latrans) body weights are variable. 
They typically weigh 8.2 - 13.6 kg in the western part 
of their range, about 13.6 kg in the Midwest United 
States, and 15.9 — 18.2 kg in the northeastern part of its 
range (see Parker 1995 [and sources within] and Young 
and Jackson 1951). Parker (1995) reported record 
weights of Coyotes in the northeastern United States as 
25.9 kg from Nova Scotia and 25 kg (Prince Edward 
Island) for two males and 23.4 kg (Nova Scotia) for a 
female, but this animal was suspected (based on breed- 
ing season) of being a Coyote-Dog (Coy-dog) hybrid. 
Dobie (1947) reported large specimens from California 
(30.9 kg) and Wyoming (32.3 kg). Young and Jackson 
(1951) described a 24.3 kg Coyote from Michigan and 
one weighing 34.0 kg from Wyoming, which was 160 
cm long. All Coyotes were reported as fat males (except 
for the female Coy-dog that Parker reported); however, 
reported values could be suspect because all data came 
to those authors secondhand. For example, it is not 
known if Dobie and Young and Jackson were referring 
to the same Coyote (with a different weight given from 
each source) from Wyoming. Finally, JGW (Unpub- 
lished data) hand-raised a wild-born male Coyote from 
Cape Cod, Massachusetts, that fluctuated between 
22.7 — 25 kg between the Coyote’s first and second year 
of life. The literature rarely reports Coyote weights 
exceeding 22.7 kg (50 Ib) and apparently never has for 
a female. 

The Coyote dubbed “Casper” (Way 2000 — ID# 
9804) was originally captured in a box trap (Way et al. 
2002a) in November of 1998 within the town of Barn- 
stable on Cape Cod, Massachusetts. She was a robust 
2.5 or 3.5 yr old female that weighed 23.2 kg and was 
148 cm long (tip of nose to tailtip) at the time. She was 
recaptured on 6 March 1999 and released without being 
handled. On 26 July 1999 she was recaptured and re- 
collared and weighed 19.6 kg and was 140 cm long. 
She was visibly skinny having raised > 4 pups that 
summer (Way et al. 2001). She tested negative for 
heartworm (L. Venezia, Hyannis Animal Hospital, 
unpublished data). On 11 March 2004 we recaptured 


9804 ca. 0.5 km from her original capture location (and 
in the core part of her territory: Way et al. 2002b) and 
replaced her old radio-collar (the battery had been dead 
for two years). She weighed (on a digitally calibrated 
scale) 25.1 kg (55.3 Ib) and was 157 cm long (body 
length [nosetip to tailbone] = 108 cm). She was robust 
with plenty of fat (1.e., her hip bones were difficult to 
palpate), her stomach had ~ 2.3 kg of meat scraps (i.e., 
bait used to capture her), and she was 4 - 5 weeks preg- 
nant, based on a physical exam and her prior denning 
habits (Way et al. 2001). Her canine teeth (0.8 — 1.2 cm) 
were worn to about half their original length. Despite 
her noted vigor, she tested positive for Lyme’s disease 
and heartworm (L. Venezia, unpublished data). Never- 
theless she had been the breeding female of a pack of 
> 3 —4 Coyotes (not including pups) for = 6 years (Way 
et al. 2002b). 

Coyotes from the New England region are large 
(Silver and Silver 1969; Wayne and Lehman 1992) and 
various theories for this have been postulated (Thurber 
and Peterson 1992) with hybridization from a small 
species of Wolf (Cais /vcaon) a very plausible reason 
(Gompper 2002; Wilson et al. 2003). Coyote #9804 
could not be a Coy-dog because she consistently gave 
birth in early April which is when wild canids normally 
whelp (Parker 1995; Way et al. 2001); Coy-dogs have 
a phase shifted reproductive cycle and typically whelp 
2 —3 months prior to wild coyotes (in mid-winter: Men- 
gel 1971). Coyote #9804’s blood is currently being ana- 
lyzed (B. White, Trent University, unpublished data) to 
determine the genetic makeup of this unusually large 
sized Eastern Coyote. 


Acknowledgments 

This study would not have been possible without 
the support from L. Venezia and his staff at the Hyannis 
Animal Hospital, equipment purchases from E. Strauss 
and Boston College, Green Grant Youth Council sup- 
port, and in-kind donations from the Way family. Two 
anonymous reviewers provided helpful comments on 
the manuscript. 


140 


Literature Cited 

Dobie, J. F. 1947. The voice of the coyote. University of 
Nebraska Press, Lincoln, Nebraska. 

Gompper, M. E. 2002. Top carnivores in the suburbs? Ecolog- 
ical and conservation issues raised by colonization of north- 
eastern North America by coyotes. BioScience 52: 185-190. 

Mengel, R. M. 1971. A study of dog-coyote hybrids and 
implication concerning hybridization in Canis. Journal of 
Mammalogy 52: 316-336. 

Parker, G. R. 1995. Eastern coyote: the story of its success. 
Nimbus Publishing, Halifax, Nova Scotia, Canada. 

Silver, HL, and W. T. Silver. 1969. Growth and behavior of the 
coyote-like canid of northern New England with observa- 
tions on canid hybrids. Wildlife Monographs 17: 1-41. 

Thurber, J. M., and R. O. Peterson. 1991. Changes in body 
size associated with range expansion in the coyote (Cais 
/atrans). Journal of Mammalogy 72: 750-755. 

Way, J. G. 2000. Ecology of Cape Cod Coyotes (Canis 
/atrans var.). M.S. thesis, University of Connecticut, Storrs. 
107 pages. 

Way, J. G, P. J. Auger, I. M. Ortega, and E. G. Strauss. 
2001. Eastern coyote denning behavior in an anthropogenic 
environment. Northeast Wildlife 56: 18-30. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Way, J. G., I. M. Ortega, P. J. Auger, and E. G. Strauss. 
2002a. Box-trapping eastern coyotes in southeastern Massa- 
chusetts. Wildlife Society Bulletin 30: 695-702. 

Way, J. G., I. M. Ortega, and P. J. Auger. 2002b. Eastern 
coyote home range, territoriality and sociality on urban- 
ized Cape Cod, Massachusetts. Northeast Wildlife 57: 1- 
18. 

Wayne, R. K., and N. Lehman. 1992. Mitochondrial DNA 
analysis of the eastern coyote: origins and hybridization. 
Pages 9-22 in Ecology and management of the eastern coy- 
ote. Edited by A. H. Boer, Wildlife Research Unit, Univer- 
sity of New Brunswick, Fredericton, New Brunswick, 
Canada. 194 pages. 

Wilson, P. J., S. Grewal, T. McFadden, R. C. Chambers, 
and B. N. White. 2003. Mitochondrial DNA extracted from 
eastern North American wolves killed in the 1800s is not 
of gray wolf origin. Canadian Journal of Zoology 81: 
936-940 

Young, S. P., and H. H. T. Jackson. 1951. The clever coyote. 
The Stackpole Company, Harrisburg, Pennsylvania. 


Received 10 May 2004 
Accepted 28 February 2005 


Book Reviews 


ZOOLOGY 


Amphibiens et reptiles du Quebec et les maritimes 


By Jean-Francois Desroches et David Rodrigue. 2004. Editions 
Michel Quinton, Waterloo, Quebec JOE 2NO. 288 pages. 
Can $40.00. 


The eye of the reader is quickly overwhelmed by 
blaze of sharp, high quality colour photographs through- 
out virtually every page of the body of the text, pages 
11 to 235. These are the combined work of 20 pho- 
tographers, with the authors contributing the majority, 
DR credited with 97 and J-FD with 42 compared to 
103 by all others. This section covers preface, forward, 
introduction, definitions of amphibian and reptile, a 
code of ethics for herpetologists, observation and con- 
servation of amphibians and reptiles, introduction to 
each major grouping within the two classes and indi- 
vidual species accounts. The latter include description, 
similar species, reproduction, habitat, critical area, food, 
predators, status, notes. These accounts demonstrate 
the authors’ familiarity with the varied characteristics 
and ecology of each species. 

A map covering from the western border of Ontario 
to the Maritimes and the adjacent northern United States 
depicts each species range in a splash of red. Those for 
the Quebec portion are drawn from the comprehen- 
sive database of the Quebec Atlas for which Rodrique 
is now the chief coordinator. This project is based at 
the St. Lawrence Valley Natural History Society and 
has financial supported from la Societe de la faune et 
de la parcs du Quebec. The data for the Maritime maps 
was provided by Don McAlpine at the New Bruns- 
wick Museum. The only problem with these is that 
the maps are too small allow detail and disjunct pop- 
ulations are easily overlooked. 

In all, 40 species are given detailed treatment (21 
amphibians and 19 reptiles (nearly half of the species 
total for Canada). Two sea turtles are not recorded for 
Quebec and two frog species, two freshwater turtles 
and two snakes are absent from the Maritimes. An ad- 
ditional account covers the complex subject of diploid 
and polyploid salamanders involving hybridization 
between species of the genus Amystoma. Three more 


Mammals of Australia 


By J. Turner. 2004. Pensoft Publishers. Acad. G. Bonchev Str., 
Bl. 6, 1113 Sofia, Bulgaria. 216 pages. Eur 38.50 Cloth. 


Having a particular interest in camels at this time, 
the first thing I tried was to look up this species. The 
book is arranged alphabetically by common name. So 


reptiles are either introduced (Red-eared Slider turtle, 
Eastern Box Turtle) or reported but not confirmed 
(Timber Rattlesnake). 

Pages 236 to 288 conclude the book. These include 
diagrams of key features and identification keys to adults 
of both amphibians and reptiles, a standard in many 
guides. But in addition, there are keys to amphibian 
eggs, salamander larvae, tadpoles (frog larvae), and cast 
skins of snakes, subjects often poorly covered or omit- 
ted in other guides. Conservation is emphasized with a 
tables presenting the number of individuals for each 
of the nine common native species which can be held 
without permit in captivity in Quebec (two salamanders, 
toad, five frogs and the garter snake). Another table 
gives the status assigned to species whose continued sur- 
vival is of most concern in Quebec, New Brunswick, 
and Nova Scotia. Groups coordinating observations of 
amphibians and reptiles in each of these provinces are 
also listed with addresses. Additional appendices cover 
the etymology of all scientific names and a glossary 
of 100 technical terms from “aire de repartition ou de 
distribution” to “zooplancton’’. Although this, like the 
entire text, is in French, it facilitates the reading of the 
book for anyone with at least a rudimentary grasp of 
the language. 

The bibliography is arranged by guides and books, 
journal articles, and technical reports. The latter is par- 
ticularly valuable as it includes work not widely refer- 
enced elsewhere. The Quebec literature is particularly 
comprehensive but the Maritime references are more 
sparse due to the greater familiarity of the authors with 
the literature of their native Quebec. 

The authors deserve the highest praise for their con- 
ception, enthusiasm, care, and toil to bring this guide to 
completion, as does the publisher for the suburb quality 
of the production. It is the new benchmark for other 
provinces to try and match. 


FRANCIS R. COOK 


Researcher Emeritus, Canadian Museum of Nature, Ottawa, 
Ontario K1P 6P4 Canada. 


I tried “Camel” without success. Next I looked under 
D for “Dromedary”, also without success. By going 
through page by page I found “One-humped Camel” 
under O. Under this system I noted that House Mouse 
and Humpbacked Whale shared a page — an odd com- 


141 


142 


bination! The index is arranged in exactly the same way 
so is no help and is superfluous. This was not an impres- 
sive start. There is a “Quick Index” arranged in taxo- 
nomic order that is helpful if you know the scientific 
names and sequence of mammalian orders. 

The information on the One-humped Camel (and the 
House Mouse, Humpbacked Whale etc.) was clear, 
meaningful and concise. Camels are now wild in Aus- 
tralia and these feral residents probably represent the 
most “natural” population in the world today. Certainly 
they seem to be extinct in their land of origin (America) 
and their classical home in North Africa-Arabia. 

This arrangement by common name renders it dif- 
ficult to make comparisons. The common kangaroos are 
under E, W and R (Eastern Grey Kangaroo, Western 
Grey Kangaroo and Red Kangaroo) You have to flip 
between widely separated pages to find the differences 
in morphology and range. There are places where you 
could see all three species in the same location (such 
as Northwestern New South Wales.) and a direct eval- 
uation would be useful. 

The range maps are a little confusing. A typical map 
shows the range in blue. Other maps are coloured light 
green. At first I thought this colour was used to show 
the range of the extinct species shown in the book. Later 
this was less clear as it was also used for subspecies 
and endangered species. I searched for an explanation 
of the range maps but did not find one. One curiosity 
I noted with the range maps is that only three creatures 
are continent-wide — the Echidna and two introduced 
species, the Cat and House Mouse. (Five others are al- 


Birds of the Raincoast: Habitats and Habitat 


By H. Thommasen and K. Hutchings, with R. W. Campbell and 
M. Hume. 2004. Habour Publishing, PO Box 219, Madeira 
Park, British Columbia, VON 2HO 222 pages, Can $44.95. 


This nice book describes in very delightful terms 
some components of the great Canadian biodiversity 
heritage: birds of the fascinating raincoast. It offers more 
than simply a nice coffee table book of a unique loca- 
tion and its fauna. Instead, this book consists indeed of 
a very sophisticated and well-written text covering all 
relevant aspects about birds, including species descriptions 
at given habitats in the stunning Canadian province of 
British Columbia. 

The text and the brilliant photos are presented by 
four writers and nine photographers. The nine chapters 
deal with birds across seasons in the various and com- 
plex habitat types of B.C.’s raincoast (south and cen- 
tral coast, but not its islands). 

This book is written by some of the bird and photo 
experts in the province. It cannot be denied that the 
traditional Anglophone, if not English, culture domi- 
nates in this book, as this culture also somewhat dom- 
inates the B.C. bird scene. No wonder, the first author 
is a country doctor, the second author makes a living 
in an English Department. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


most continent-wide, Common Walleroo, Gould’s Wat- 
tled Bat, Red Kangaroo, Rabbit and Dingo) Many of the 
rest are quite localized. 

The illustrations are odd. Eventually I realized they 
reminded me of children’s tracings. The shape is most- 
ly correct but gets smoothed out in places. The Red Fox 
is an odd looking creature. The accuracy of the colours 
varies. For example the Mountain Possum is grey not 
brown, but I think the pointed tail make the identifica- 
tion clear. The Greater gilder is depicted as grey- 
headed and white-tailed when it can be the reverse. 
The text somewhat clarifies this distinction. The odd- 
est colour rendering of all is the silvery grey Antarctic 
Fur Seal. All those I have seen are rich brown. However, 
some of the illustrations are quite good. The Lead- 
beater’s Opossum , the Agile Wallaby and the Banded 
Hare Wallaby all seem to be fairly accurate. 

Despite my criticisms I do not think anyone will have 
a problem identifying a mammal though. Using the text, 
range map and the illustration should enable you to 
reach a clear conclusion on any of the 319 species cov- 
ered. (This is a little short of the accepted total of 379 
and I think those missing are some bats and marsupials, 
plus most of the cetaceans.) So, as there is sufficient 
diversity and differences in range you can still use this 
book as a field guide. The size (21 x 28.5 cm or 8.25 
x 11 in.), however, means it is not truly portable. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 
K1J 6K5 Canada. 


The bird photos are among the best I have ever seen 
for B.C. Some readers might find bird photos taken at 
the nest old-fashioned, and in some circles they deemed 
to be unethical even (since disturbing birds on nests and 
thus not publishable). 

The photo layout and arrangement of the text strikes 
a great balance between detail and text flow; for the 
bird enthusiast this book is simply a joy to read and to 
browse through. Popular bird names are explained, and 
the poetry literature link is made by citing Anglophone 
poets from U.S., Canada and UK, such as Archibald 
Langman, William Blake, John Burroughs, Duncan 
Campbell Scott, John Clare and others. Personally, I 
specifically like the raptor, owl and sparrow sections, 
and the photos of the wood-warblers, blackbirds, owls, 
flowers and plants. 

The wording in this book is extremely careful. How- 
ever, I do not agree with the statement made by the 
authors that long-distance migration 1s a risky business 
for birds. Some species actually living that life-style 
have the longest life span known for birds; e.g., Terns 
and Godwits. . 

Allowing for a more global context, I really like 
that wintering grounds for B.C. birds are named as well, 


143 


which can wander off to Nicaragua and other tropical 
locations. However, some more spatially explicit infor- 
mation would be helpful for people exploring B.C. and 
its birds, too; e.g., a map and location names where 
bird species can be found in B.C. The authors clearly 
must know these details but so far don’t share them 
with the reader. 

Hardcore birders might be happy to learn that the 
book addresses how to avoid confusing Mallards from 
Shoveler, but no distinguishing field photos are shown 
really for identification headaches such as Common 
and Barrow goldeneyes, the species of mergansers, 
loons, grebes and crows. Issues of rare birds are basical- 
ly ignored. So one will not learn about Eurasian strag- 
glers from Russia that excite the B.C. birding commu- 
nity much and which increase birding tick lists. Avian 
subspecies do not get covered 1n depth, but some avian 
genetic issues are addressed. 

Fascinating details on birds in B.C. get shared: The 
first Barred Owls were seen on the B.C. coast in 1966. 
The re-use of nests from owls and cavities by other birds 
are explained, the reader learns what “Sap Wells” are, 
and what makes for dabbling and diving ducks. Winter 
survival strategies are explained, the concept of Nest- 
Helpers is introduced, as well as the odd behavior that 
Rosy-finches cover the nest with their wings. I also like 
the mentioning of (sea) mammals, and how glaciation 
provides the reason that Myrtle and Audubon warblers, 
two waxwing species, and Red- and Yellow-shafted 
flickers can be found in this huge province, which is at 
least two times larger than California. The authors 
mention that some people in B.C. perceive birds as 
pests; e.g., Golden-crowned Sparrows. The Bird Check- 
list and the Index are nice additions making this book 
a well structured resource. 

The authors cite John Clare: “To study birds without 
reference to their habitats is to examine their carcasses 
in glass cases”. And here I find the book a little weak. 
Major issues for B.C., and which every bird enthusiast 
will notice in no time during an outdoors trip, such as 
habitat loss, pollution, urbanization and boating and 
their excessive human foot prints, etc. are not sufficient- 
ly mentioned when considering its magnitude and im- 
portance for B.C. Environmental buzz words such as 
clearcuts, fragmentation, oil spills or even Clayoquot 
Sound and Carmanah Valley do not really exist, nor is 
there a mention of Fraser River, or Queen Charlotte 
Islands and pelagic habitats. At least it gets mentioned 
that Glaucous-winged Gulls have increased three times 
in B.C. during the last 5O years (no reasons given), 
and that many river populations of Euchalon have been 
lost. Changes in Cormorant and Raven numbers and 
other related information are not mentioned. 

This book might not fully satisfy readers that have a 
strong science hunger: Scientific names are not present- 
ed, researchers are rarely named (except for Margaret 
Nice and her Song Sparrow work) and this books lacks 
quantitative statements, other than a given home range 
of the Pileated Woodpecker (1000 acres) and that the 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Brown-headed Cowbird population would run up to 
50 million individuals. Initially, I found that the authors 
overemphasize a little that birds would be monoga- 
mous, but the Extra-pair Copulation (EPC) issue gets 
mentioned in the text later. [ cannot agree with the 
impression somewhat portrayed in the book that feed- 
ing birds in winter would do any good to them, and 
that birds would not survive otherwise. Birds actually 
evolved fairly well without bird feeders and did well 
without any human intervention over the last hundred 
thousands of years. 

If one wants to be really picky, one can have a little 
seabird issue with this book though. Short-tailed Alba- 
tross, Cassins Auklets and Tufted Puffins are not men- 
tioned. The massive Old-growth Forest habitat loss 
relevant for Marbled Murrelets get only mentioned 
marginally. I am quite sure that the 8500 figure, pre- 
sented by the authors as the total British Columbia 
breeding population for Marbled Murrelets is a drastic 
underestimate. The latest overall population estimate 
for B.C. is 60000 and some models suggest even over 
150 000 birds (this includes non-nesting birds). There 
is no Marbled Murrelet nest photo presented; a key 
icon for how birds can serve as habitat indicators for 
the raincoast. The Bird Checklist is nice but does not 
fully match my experience: Marbled Murrelets for 
instance are not rare, but can in some fjords and during 
winter be among the most abundant species in coastal 
waters even (as for instance the case in Clayoquot and 
Desolation Sound). Despite all this great detail, existing 
expert knowledge of the authors and a wonderfully 
detailed text, I am not clear why the first Canadian 
Marbled Murrelet nest found in 1992 by Volker Bahn 
and others is not mentioned (but the first one found 
1974 in California gets presented). 

One point that puzzles me, and which might explain 
some issues I would summarize from above as being 
“uncritical” or “unaware” about environmental rain- 
coast issues in B.C., is a phrase in the acknowledgments: 
“Weyerhaeuser and BC Hydro served as a source of 
inspiration and researching this book”. Most environ- 
mentalists and bird enthusiasts actually might not relate 
these major companies with inspiration, nor perceive 
them as a great guide for research. 

Despite its potentially huge value, I am also puzzled 
by a text feature presented in the appendix dealing with 
“Proactive Conservation in British Columbia: The Wild- 
life Data Center’. In recent years, much of British Co- 
lumbia’s conservation has been severely affected, if 
not harmed, by the lack of free and shared high quality 
data among experts and agencies. The professional con- 
servation climate in British Columbia is actually quite 
“polluted” due to this situation. Taken from its des- 
cription, this data center holds much of the urgently 
needed information which is so crucial to sustainable 
and progressive land and habitat management and con- 
servation of wildlife and birds in the province. Des- 
pite its claim to be pro-active no website is presented 
to the audience. I miss the word “digital” in there, too. 


144 


Here is where the cultures and generations truly clash. 
And how does this relate, link and compare with what 
the Canadian and provincial government does (see for 
instance Geogratis website http://geogratis.cgdi.gc.ca/)? 
Here a change and update to the new millennium is 
needed for British Columbia so that high quality bird 
and conservation data are freely shared and made avail- 
able to the global public over the internet nowadays. 
The description of the data center mentions for instance 
that they hold the largest nest record data pool for 
Canada with 180 000 records! Let’s put these massive 
data sets on the public table for much of the urgently 
needed Conservation Management in British Columbia. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


The book jacket reads: “Perfect for Birdwatchers, 
Naturalists and Environmentalists.” 

The reader might decide him- or herself on the phil- 
osophical question how much birds can and should be 
used to address environmental concerns, and whether 
they contribute to conservation. I recommend this book 
for sure as a very nicely written description and pho- 
tographic explanation of birds in British Columbia, as 
well as a celebration of a Canadian and world heritage 
component. 

FALK HUETTMANN 


Institute of Arctic Biology, Biology and Wildlife Department, 
University of Alaska-Fairbanks, Fairbanks, Alaska 99775 
USA 


The Bird Almanac: A Guide to Essential Facts and Figures of the World’s Birds 


By David M. Bird. 2004. Key Porter Books Ltd. 6 Adelaide 
Street 10 Floor, Toronto, Ontario, M5C 1H6. 460 
pages, Can $24.95 Paper. 


Within this book is a massive amount of bird-related 
information! This review could actually end right here. 
However, I will elaborate somewhat. 

Information in this book is presented in three ways: 
line drawings (restricted to the anatomy chapter), tables/ 
lists as well as glossary-style entries. These are all ap- 
propriate and lead the reader to finding information 
rather quickly. There is both 4/7@img information and 
ornithological information (and a massive amount of 
overlap which is shared by both). A glossary of over 
20 pages contains over 1000 terms — surely the word 
you’re looking for must be there! 

There are simply too many categories of informa- 
tion to give more than a smattering of examples here. 
Both traditional and genetically-based classifications 
of bird families are given, followed by the massive list 


Common Birds of Ontario 


By J. Duane Sept. 2004. Calypso Publishing, P.O. Box 1141, 
Sechelt, British Columbia, VON 3A0. 94 pages, Can. $12.95 
Paper. 


This slim volume is an entry in the “common birds 
to know” category. It presents some 142 species, most- 
ly two per page, each with a small coloured photograph 
of the bird, together with a brief description, some 
information on size, nesting and habitat, followed by 
a paragraph or two on some topic of more general 
interest. A few accounts include a “Similar Species” 
category, and about 10% of the species receive a full- 
page treatment with two pictures. 

There’s a role for well-thought-out books of this 
kind. Persons who are mildly interested in birds, but 
who are not ready for a comprehensive guide, might 
skim through the pages of a book of this kind, and try 
to match up the bird they had just seen with one of the 
illustrations. But to be really useful, the author must 
give careful thought to the selection of species and to 


of all known bird species. Significant people are list- 
ed in several tables, ornithological award recipients, 
world-class listers, Taverner Cup winners, bird artists 
and more. Bird watching clubs, ornithological societies, 
magazines and journals are listed from sources around 
the planet. 

To compare this book with Leahy’s 7he Birdwatch- 
er ’'s Companion, a recent tome of similar intent, would 
be to have 7he Bird Almanac come on top. The former 
is essentially all in dictionary format, and therefore lacks 
the comparative ease (or the ease of comparisons) of 
the thematically-organized, tabular format in A/manac. 
There are more in-depth definitions and descriptions 
in Companion (it is also a much bigger book), and the 
bibliography is better- organized, but I believe birders 
will much more evyoy flipping through A/manac. 


RANDY LAUFF 


Biology Department, St. Francis Xavier University, Antigonish, 
Nova Scotia B2G 2W5 Canada 


utilizing the limited space to the best advantage, rec- 
ognizing the user will likely be a complete novice. 

Unfortunately, the present volume does not meet 
these objectives; in fact, I was left with the feeling that 
the author himself does not know Ontario birds very 
well. Problems abound, and there is only space here for 
a brief sampling. The selection of species is puzzling: 
avocet and Wilson’s Phalarope are in but Mute Swan, 
Eastern Phoebe and grackle are missing. Lesser Scaup 
appears but Greater is not mentioned; Tennessee and 
Cape May warblers are shown but Magnolia and Black- 
and-white are not. 

Some plate selections are poor: for example, the Red- 
tailed Hawk appears to be of the western race and could 
confuse a beginner; and the Least Flycatcher’s wingbars 
are almost wholly concealed. The text is also misleading 
in places. We’re told wigeon are “often” found winter- 
ing in large flocks on golf courses, Long-tailed Duck is 
a “common migrant’, but “not often seen in large num- 


145 


bers’, Gray Jays are widespread and Wilson’s Warblers 
are “common in moist areas”. The foregoing are just a 
few examples of inaccuracies which could have been 
readily avoided, and which are bound to mislead the 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


potential reader. All in all, this is not a book I can rec- 


ommend. 
CLIVE E. GOODWIN 


1 Queen Street Suite 405, Cobourg, Ontario K9A 1M8 Canada 


Frogs of Australia: An Introduction to Their Classification, Biology and Distribution 


By James R. Turner. 2004, Pensoft, Sofia-Moscow. 163 pages. 
Price not available. 


Australia is renowned for its mammals, unique as a 
consequence of that continent’s long separation from 
the other land masses. Of the amphibians, only frogs 
are represented; both the caecilians of other southern 
continents and the salamanders characteristic of north- 
ern continents, are absent. 

Frogs of Australia presents 218 forms (213 species, 
one divided into 5 subspecies and one into 2); 110 of 
these named since 1960. The five families represented 
contain 29 genera. Three families are held in common 
with Canada. Hylidae occurs in the Americas, Europe, 
Asia and northern Africa but its 76 Australia species 
are often separated as the Pelodryadidae). Bufonidae 
is represented only by the Cane Toad, 4u/fo marinus, 
introduced from Hawaii in 1935 which, despite its near 
legendary reputation for decimating the native Aus- 
tralian frog and small mammal fauna, has spread only 
in the eastern edge of the continent, occurring in the 
Northern Territory, Queensland, and New South Wales. 
Ranidae, of the Americas, Eurasia and Africa, occurs 
only marginally with a single species only in the Cape 
York Peninsula in the northeast. Surprisingly, the latter 
resembles our Canadian Wood Frog, Rana sy/vatica, 
in pattern (dark mask) and call (“like the sound of a 
duck but lacks a musical quality”’). Of the other three 
families, two are not represented in Canada: the Myo- 
batrachidae 118 species (formerly included in the 
southern western hemisphere Leptobatrachidae) and 
Microhylidae 18 species (widely distributed in south- 
ern western hemisphere, Africa and Asia). 

Most forms are given a vertical half page column, 
but six have two columns (two with consecutive half 
pages; four on one entire page each). A problem for 
the non-Australian (and perhaps even some Australians) 
is that the forms are sequenced in alphabetical order 
by English names, rather than being grouped by fam- 
ily or genus. Thus, in one case two subspecies with 
distinctive common names are separated by 100 pages. 


The only other subspecies fortunately have a base com- 
mon name, each form with a modifier, so all five occur 
consecutively. 

Every form is illustrated in colour and accompa- 
nied by a range map. The text accounts are very brief: 
Latin name, Synonyms, Family name, Habitat, Distri- 
bution, Length, Abundance, Status, Meaning (of Latin 
name), Behaviour (generally breeding and call), Devel- 
opment (number of eggs, where laid), and Locality 
(political divisions it occurs in). Although all are by 
the author, the illustrations vary from superb to less so, 
with those in which a bit of substrate is included more 
deftly done while the others give the effect of crude 
cutouts. But as Dr. T. J. Hawkeswood points out in the 
Forward they “add a 19" century feel to the book”. No 
variations are depicted. The maps apparently give a 
background of regional elevation in dark green, light 
green and tan though this seem not to be explained any- 
where and the colours could be taken for vegetation 
zones. Superimposed, the purple depiction of range 
effectively stands out for all but the most restricted dis- 
tributions. The superb layout facilities flipping through 
for an overall impression of variation and ranges. 

Additional text is minimal. A Quick Find Index lists 
families and contained species by scientific name and 
page number of account, and an Introduction covers top- 
ics such as How Old Are Frogs (at least 180 million 
years), Common and Scientific names, Species and 
subspecies, descriptions, distribution. A page of dia- 
grams depicts Diagnostic Characters of Frogs, four 
pages list the characters of the families and genera 
with comments on conservation. An outline map gives 
the eight political divisions and a two or three letter 
code for each (but West and South Australia are both 
designated WA whereas the latter is properly SA in the 
text). The book concludes with a two-page Glossary 
(address to xeric) a 20 pages of references and a three- 
page index to common and scientific names. 


FRANCIS R. COOK 


Researcher Emeritus, Canadian Museum of Nature, Ottawa 
K1P 6P4 Canada 


Experimental Approaches to Conservation Biology 


By Malcolm S. Gordon and Soraya M. Bartol. 2004. Uni- 
versity of California Press. Berkley, California. xv + 343 
pages, $75 US. 


The papers collected together in this volume were 
originally prepared for a conference of the same name 
held in 2001. The editors have brought together re- 
search that emphasizes both basic and applied scien- 


tific experiments related to conservation, particularly 
those at the organism level. 

The book consists of four sections. The first section, 
Introduction, consists merely of a five-page introduc- 
tion along with a one-page list of conservation-related 
websites. The list of web resources is very basic and 
is limited to five government sites (one Canadian and 
four American) along with a number of major NGOs 


146 


such as the Nature Conservancy and World Wildlife 
Fund. Each of the remaining sections begins with an 
overview of the section and then papers on that topic. 
The second part of the book consists of seven papers 
on the theme of “Conservation of Endangered Species.” 
The papers cover a variety of species (e.g., tree snails, 
rock iguanas, New Zealand birds, Australian marsupials) 
and topics (e.g., er s/f propagation, amphibian decline, 
conservation endocrinology). Section three, “Control 
or Elimination of Exotic and Intensive Species” fea- 
tures three papers on the subjects of California grass- 
lands, tree invasions and the use of biological controls 
on exotic species. The final section of the book, “Policy- 
related Matters” contains four papers. Topics include 
the case study of the US Army and the Desert Tortoise 
and the challenge of biodiversity conservation within 
African national parks. 

As is usually the case in such volumes, the papers 
vary substantially in their scope and quality. At one 


Gulls of North America, Europe, and Asia 


By Klaus Malling Olsen and Hans Larsson. 2004 $55.00 Cloth. 


My reaction on opening this book was akin to the 
one I had when I first saw Seabirds! — this is a giant leap 
forward. It is a glorious wad of text accompanied color 
paintings (by Larsson), wonderful range maps and a 
multitude of color photographs. The authors say they 
cover forty-three species of gull. I found there are 50 
species of gull worldwide that have generally been 
accepted in the recent literature. Of the 43 covered in 
this book 37 are on the world list. The remaining species 
are splits made and justified by the authors. These are 
Thayer’s, American Herring. Caspian, Vega, Heuglin’s 
and Common Gulls. Adding these in would bring the 
world total to 56. Also included are three South Ameri- 
can vagrant species that are covered by a short text and 
a couple of photos. This means the book covers three- 
quarters of the world’s gull species. 

Each species is described in extensive detail, with a 
full description of all plumages at all life stages and 
racial variations. The accompanying illustrations sup- 
port the text in these details. Additionally there are some 
comparison plates comparing similar species. 

So this is a book of detail and it is in the detail we 
find the devil. Gull taxonomy is notoriously complex, 
poorly understood and subject to argument. I decided 
to examine first the Herring Gull and the newly-split 
American Herring Gull. Comparing the descriptions 
with the author’s illustrations of the adult, I found it 
difficult to discern the majority of the “differences.” I 
have noted similar bird-to-bird variations in single flocks 
of each one of these “species.” I quickly realized I was 
concentrating on the photos and not the paintings, as I 
found the depictions too pale. The photos were closer 
to the pearl grey of my memory. There are comparison 
plates of large gull taxa (a good idea) where the Amer- 
ican Herring Gull is shown as paler than an Iceland Gull. 
This is not an error as it is set in the middle of Glaucous 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


extreme, Michael Hadfield and collaborators provide 
an overview of the over 20 years of work his lab has 
done on the conservation of Hawaiian tree snails of the 
genus Achatinella. Their work has combined mark- 
recapture field studies, devising field exclosures to 
protect populations, developing protocols to maintain 
populations in the lab, and developing conservation 
priorities through the use of genetic markers. In con- 
trast, other papers focus on a single issue or experiment. 
In addition, although all the papers were prepared for 
this book they vary in format, with only some of the 
papers including a summary. Nonetheless, this is a 
strong collection of papers, displaying the breadth of 
experimental conservation biology. 


DAVID SEBURN 


Seburn Ecological Services, 2710 Claenda Street, Ottawa, 
Ontario K2B 7S5 Canada 


and Glaucous-winged Gulls, and not with the European 
Herring and Yellow-legged Gulls. 

The comparison of the non-adult plumages is even 
more difficult. Picking your way through the myriad of 
details of multiple plumages for each species is time 
consuming and can be tedious. This is not a criticism 
nor a question of right and wrong, but more the part of 
an ongoing debate. It has prompted me to look more 
carefully at the local American Herring Gulls this year. 

Nowhere does this taxonomic debate emerge more 
clearly than with Thayer’s Gull. The author notes “Tax- 
onomic status not fully clarified”’ Many years ago I 
went through three trays of study skins at the Cana- 
dian Museum of Nature in Ottawa. The first tray con- 
tained Iceland gulls. The second contained Thayer’s 
Gulls. The problem was the third tray had birds that 
looked like one of these species on their left side and 
the other species from the right side. In addition, during 
my years in Nova Scotia the commonest winter bird 
in my backyard was the Iceland Gull and I soon real- 
ized that it was a very variable species. 

Next I turned my attention to the adult Great Black- 
backed Gull. This raised further questions. The text says 
“Legs flesh, sometimes with a grey or creamy (rarely 
yellow or extremely orange) Age.’ (my emphasis). Legs 
flesh, sometimes with a grey or creamy tinge certainly 
fits the birds I have seen. I have found a yellow-legged 
bird. The next species is the Kelp, also large and 
black-backed. The adult’s feet are yellow but could be 
greyish-olive or greenish. When I, carefully (you do 
not want to damage the fine photos), curl the photos 
back so I can see the two birds side by side they are 
very similar, separated by their feet colour. If we had 
a grayish-footed Kelp Gull wander north would we 
notice it and would I get excited at a yellow footed 
Black-back? Yellow legs is the classic mark for separat- 
ing the Lesser Black-back Gull. 


147 


Clearly identifying gulls is fraught with lots of pitfalls 
for the less than careful. Identification of vagrant gulls 
need to be done with vast care and all the field marks 
need to be examined. Likewise we should not pass off 
our local crowd of gulls as being the usual species; a 
great treasure could be hidden in the flock. 

The pros and cons of gull taxonomy notwithstand- 
ing, this book is a wealth of information. The same 
exacting detail is given for all the species in the book, 
making it a treasure trove for the dedicated birder. 
Regardless of your opinion on the status of the species 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


splits, there is much material for good scientific dis- 
cussion and a basis for careful field observations. This 
all leads to the final question — why not go all the way 
and write Gulls of the World. After all it is only 13 more 
species, barring splits! 

Roy JOHN 
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 

K1J 6K5 Canada 

Literature Cited 


Harrison, Peter. 1983. Seabirds: An Identification Guide. Helm Field 
Guides Houghton Mifflin Co., Boston, Massachusetts 


The Monarch Butterfly: Biology and Conservation 


By Karen S. Oberhauser and Michelle J. Solensky, Eds. 2004. 
Cornell University Press, New York. vil + 248 pages, illus. 
539.95 US. 


This book consists of a collection of scientific papers 
on the Monarch, divided into chapters on reproduc- 
tion, migration and overwintering biology as well as 
some miscellaneous papers on general topics. Each of 
the four chapters 1s prefaced by a brief overview writ- 
ten by the editors. Karen Oberhauser is an Assistant 
Professor at the University of Minnesota, while Michelle 
Solensky is with the Department of Biology at the 
University of St. Thomas. The information presented 
includes observations from each of the three major 
Monarch populations worldwide (eastern and western 
North America and Australia). 

This book contains a great deal of interesting infor- 
mation, presented as original scientific literature. The 
majority of the papers are well-written, although some 
are likely to be overly technical (especially in terms of 
the statistical analyses) for some readers. Of particular 
interest are the papers on citizen-based research pro- 
grams, which present data collected by widespread 
networks of volunteers. These papers demonstrate how, 
with relatively modest training and logistical support, 
public participation can be used to enable research 
into wide-ranging populations which could otherwise 
prove difficult or impossible to monitor effectively. 
The overviews which introduce each of the four chap- 


ters help to make the information presented in the book 
more accessible to a broader audience (including, per- 
haps, some of the volunteers whose efforts are reported). 

The black-and-white photographs which accompany 
each overview are excellent. However, the overall pau- 
city of illustrations (other than the numerous graphs 
contained within the various papers) is regrettable. With 
such a colourful and attractive subject as the Monarch, 
surely it would not have been difficult to find more 
photographs to include in the book? Aside from their 
purely aesthetic value, additional photographs or draw- 
ings could have been used to enhance the text by illus- 
trating some of the features, behaviours and method- 
ologies described (e.g., growth and development, wing 
damage caused by courtship, experimental equipment 
design, etc.). 

In short, this book presents interesting data on every 
major aspect of Monarch biology, with additional em- 
phasis on the need to extend and incorporate such data 
into ongoing and future conservation efforts. However, 
the scientific format of the text may deter some readers, 
which is unfortunate considering the widespread public 
interest (as acknowledged in the book) in this familiar 
and beloved butterfly species. 


AMY MACPHERSON 


Jacques Whitford Limited, 2781 Lancaster, Ottawa, Ontario 
K19 1A7 Canada 


The Behavior and Ecology of Pacific Salmon and Trout 


By Thomas P. Quinn. 2005. American Fisheries Society in 
Association with the University of British Columbia Press, 
Vancouver, Canada. 378 pages, illus. Cloth CDN $85.00; 
paper CDN $44.95. 


The vast amount of literature on the Pacific salmon 
is rather intimidating. Indeed, several years ago when 
I first began conducting research on this group of fish 
after leaving the Midwest, it was difficult to know where 
to begin. Today there is a clear starting point I would 
read Zhe Behavior and Ecology of Pacific Salmon and 
Trout by Thomas Quinn. This book is an excellent 
resource for students and fisheries scientists, providing 
a broad overview of Pacific salmon ecology. However, 


this book also has broader appeal to animal ecologists 
and naturalists. Thomas Quinn is a prominent ecolo- 
gist (Professor at the University of Washington) who 
has devoted his career to research on Pacific salmon. 
He shares his passion and experience in the first com- 
prehensive book to focus on the ecology and behav- 
iour of the Pacific salmon. This book is complimenta- 
ry to other compilations on this group of fishes such 
as the Physiological Ecology of Pacific Salmon by 
Groot, Margolis, and Clarke. 

The book is focused on the salmon and trout in the 
Genus Oncorhynchus. This multi-species approach is 
one that could easily lead to confusion. However, Quinn 


148 


has provided the necessary context and background 
on basic life-history/natural history so readers, even if 
unfamiliar with Pacific salmon, will be able to follow. 
In fact, throughout the book he contrasts and compares 
interesting characteristics among species (and among 
populations) to reveal their incredible diversity. Quinn 
has also provided enough background on techniques 
in fisheries science that readers can gain an apprecia- 
tion for the challenges in answering some questions. 
For example, in the migration chapter he discusses 
techniques for monitoring individual behaviour (e.g., 
conventional tagging, acoustic telemetry). For most 
trained fisheries scientists, some of the background 
material will be superfluous. However, it is necessary 
considering the broad audience. In the end, I believe 
that even the most experienced fisheries scientists will 
find the writing style and content to be refreshing and 
not too simplistic. 

The book includes coverage of some rather novel 
topics within the 19 chapters. For example, this is the 
first fish ecology book that I have encountered that in- 
cludes a chapter on the ecology of dead fish. Several 
years ago such a chapter would have been rather short. 
However, the explosion of research in the past few years 
on the role of salmon carcasses in the environment, 
much of which has been conducted by Quinn and his 
associates, enables such a chapter to be developed. This 
chapter in particular will be of interest to more gener- 
al ecologists and naturalists. The chapters associated 
with adult migration and reproduction (2 through 6) are 
also exceptionally strong coincident with the authors 
expertise in those areas. That said, there were no chap- 
ters that I would consider weak, with my preference for 
some perhaps reflecting my personal interests and biases. 

The text is well referenced but not to the extent that 
one feels like they are wading through a scientific man- 
uscript. Quinn provides ample detail on the studies he 
summarizes, but more importantly, he provides exten- 
sive commentary and interpretation. He also poses 
questions that will serve as a springboard for countless 
graduate student projects. This is what really separates 
this book from other works that are available. His syn- 
theses draw together data from disparate studies, often 
using this information to develop original figures or 


Venomous Reptiles of the Western Hemisphere 


By Jonathan A. Campbell and William W. Lamar. 2 volumes. 
2004. Comstock Publishing Associates, Cornell University 
Press, Ithaca, New York. 870 pages. 


This is a second edition, revised and expanded, of 
the authors’ 7he Venomous Reptiles of Latin America 
published in 1989. Added to the southern species cov- 
ered in that book are those occurring in North America. 
The authors are from the University of Texas at Arling- 
ton and at Tyler, respectively. Additional contributors 
are Edmund D. Brodie II and Jr. Indiana University 
and Utah State University, Ronald L. Gutbertlet Jr. and 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


conceptual diagrams. His goal was to “inform and 
excite” the reader — a goal that I would argue he has 
more than achieved. The chapters are not exhaustive 
in coverage. Instead, Quinn has focused on selecting 
examples that contribute to an overall understanding 
of Pacific salmon and are interesting. Although his 
intent was not to advocate (as he reveals in the preface), 
the last chapter did provide some interesting final 
thoughts. As “the” authority on Pacific salmon, I would 
have appreciated if Quinn had expanded these sections 
on the current status and prognosis of Pacific salmon. 
It was this section that I found particularly fascinating, 
but it left me desiring more. 

The book is available in both hard and paperback 
form and is reasonably priced. The photograph on the 
cover is stunning (taken by the author). Other photo- 
graphs and plates within the text are equally interesting 
and of high quality. Exceptional care has been taken 
in developing crisp and clear figures that have consis- 
tent formatting throughout (e.g., axis, font, symbols). 
The author has eliminated details of most statistical 
tests presented by other authors and urges readers to 
consult original sources. Brief summaries are provided 
at the end of each chapter and the book concludes with 
an extensive index. Collectively, these characteristics 
further elevate the value of this work and contribute to 
its flow and ease of reading. 

There are few groups of fishes, or other animals for 
that matter, where one could develop a book such as this 
that would have appeal to anyone with an interest in 
animal ecology. The book is extremely engaging and 
will leave the reader with a desire to don a pair of waders 
and explore some of these topics first hand. I am con- 
fident that Quinn’s book will become a well-read and 
well-cited contribution. Shortly this book will be found 
on the office shelves of most salmon biologists, but I 
suspect it will be equally common in the personal lib- 
raries of armchair naturalists, anglers, and others with 
broad interests in ecology. 


STEVEN J. COOKE 


Centre for Applied Conservation Research, Department of 
Forest Sciences, University of British Columbia, 2424 Main 
Mall, Vancouver, British Columbia V6T 1Z4 Canada 


Michael B. Harvey University of Texas at Tyler and East 
Tennesse State University, Robert Norris Stanford 
University Medical Center, David A. Warrell, Centre for 
Tropical Medicine, University of Oxford, and Vinicius 
Xavier da Silva Universidade de Sao Paulo. 

Theirs is an accomplishment of monumental pro- 
portions in size and collation of information. It is of cof- 
fee table format and includes a staggering 1500 colour 
photographs, 135 of them pictures of snakebite effects 
about which the authors’ relate that a the first edition 
reviewer termed them their “hideous picture album”. 


149 


The species accounts begin with two lizards (the Gila 
Monster and the Mexican Beaded Lizard, genus 
Heloderma), the sole members of their family and the 
only venomous reptiles other than snakes in the world. 
Two families of snakes follow (Elapidae, including here 
the coral snakes and the sea snakes) and Viperidae (here 
represented by pitvipers) in 190 species accounts. 
Omitted are the few poisonous rear-fanged Colubridae. 
Volume | contains all species except the rattlesnakes, 
while Volume 2 covers the latter. 

The species accounts lead with scientific name and 
original describer and date, reference to figure (draw- 
ing), map (with dots of collections and shading for the 
suggested range), and plates (colour photographs), 
synonymy, local names, English name, and etymology. 
For some species a quotation from an historical obser- 
vation is added. Sections follow on distribution, habitat, 
description, similar species, and remarks (including des- 
cribed subspecies and relationships with other species). 

As comprehensive as these accounts are, there are 
other topics covered. In volume 1, there is an Introduc- 
tion, Regional Accounts and separate Keys to Canada 
and United States, Mexico and Central America, The 
Caribbean Islands, and South America. Volume 2 in- 
cludes chapters on Venomous Snake Mimicry, The 
Evolution of New World Venomous Snakes, Venom 
Poisoning by North American Reptiles, and Snakebites 
in Central and South America: Epidemiology, Clinical 
Features Management. The volume has an 11-page 
Glossary, and a 116-page Literature Cited. A 28-page 
index is repeated to conclude each volume. 

Canada has only three Cvo/a/us. The Timber Rattle- 
snake C. horridus (Ontario, now extirpated), Prairie 


Whales and Dolphins of the World 


By M. Simmonds. 2004. The MIT Press, Five Cambridge 
Centre, Cambridge Masachusetts 02142. 160 pages, U.S. 
$29.95 Cloth. 


Coffee table books of cetaceans seem to be popular 
with authors, photographers and the buying public. Here 
is another fine example. I note my local library has three 
shelves of mammal books. Almost 20% are on whales 
and the other books cover the rest of the world’s 5000 
or SO species of mammal, a testament to the populari- 
ty of cetaceans. 

While this book does not cover all the world’s species 
it does have photographs of over 50 per cent. As you 
would expect the photographs are remarkably good. All 
the common species (common that is in books) like 
Blue, Sperm Humpback, Minke are there. But so too 
are some of the less well known and photographed, 
such as the Boto and Pink River Dolphin. These latter 
animals are not so difficult to see but are stunningly 
hard to capture on film. 

The first half of the book is devoted to basic informa- 
tion on whales, similar to that found in most volumes 
of this type. The second half has information on the 
threats faced by these animals and the conservation 


THE CANADIAN FIELD-NATURALIST 


Vol... 119 


Rattlesnake C. viridis (Saskatchewan and Alberta), 
Pacific Rattlesnake C. oveganus (British Columbia) and 
one Szs7rurus, the Massasauga S. cafenatus (Ontario). 
Despite the through coverage elsewhere, the peripheral 
nature of their distributions here may account for the 
scant coverage of Canadian references the southern- 
based authors have included. Some relatively recent 
contributions by Pat Weatherhead formerly at Carleton 
University, Tony Russell at University of Calgary, and 
Pat Gregory at the University of Victoria and their many 
students are included. However, some citations for Can- 
adian distributions are merely popularized accounts by 
non-Canadians such Curran’s 1935 rattlesnake article 
in the American Museum’s Vawral History or Kozloft’s 
1976 book an animals of the Pacific northwest. Many 
are dated, the citation to the pioneering Canadian check- 
list by Logier and Toner is to the first (1955) edition 
rather than 1961 revision. Two references to Barbara 
Froom are to 1964 and 1967 articles or pamphlets 
whereas her 1972 book is omitted. The detailed distri- 
butional documentation in the Ontario Herpetofaunal 
Summary and the status reports of Committee on the 
Status of Endangered Species of Wildlife in Canada 
(COSEWIC), or the symposiums on the Massasauga 
and conservation promotional newsletter Rasilesnake 
Zales sponsored by the Metro Toronto Zoo are ignored, 
perhaps because they are not readily accessible in the 
mainstream literature. 


FRANCIS R. COOK 


Researcher Emeritus, Canadian Museum of Nature, Ottawa, 
Ontario K1P 6P4 Canada 


measures in place or necessary. I also has a very odd 
error. The author writes of the Basques from northern 
France. Any Basque will tell they always lived in the 
Pyrenees Mountains of northern Spain and southern 
France. 

The author list 83 species of whales and dolphins; 
one short of the most current list. The missing species 
is North Pacific Right Whale (Zu4a/aena japonica), 
recently separated by Rosenbaum et al, based on DNA. 
[This counting does not include De/phinus tropicalis 
as this is likely a variant of 2. capensis and not a sep- 
arate 84" species] 

Why should I buy this book? I can think of three 
reasons. Any good whale book is worth having, espe- 
cially when it such fine photos. The latter half of the 
book contains new and up-to-date information of im- 
portance. The royalties from sales are going to the 
Whale and Dolphin Conservation Society, where the 
author is Director of Science at the aforementioned 
society. 

Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 
K1J 6K5 Canada 


150 


BOTANY 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Atlas des Plantes Villages du Nanavik — Atlas of Plants of the Nunavik Villages 


By Marcel Blondeau, Claude Roy and Alain Cuerrier. 2004. 
Editions Multi Mondes, 930, rue Pouliot, Sainte-Foy, 
Quebec GIV 3N9, Canada. xxvii + 610 pages. $42.75 
(includes taxes and shipping). 


This small book (5 x 7 x 1 % inches) contains 433 
pages depicting absolutely beautiful flowering plants 
and lichens that can be found around the villages in 
Nunavik, that part of Quebec north of latitude 55°N. 
Included on each illustration page ts a distribution map 
which depicts the towns around which each species was 
found in Nunavik, together with a square mark which 
indicates its wider range into Iqualuit, Labrador, New- 
foundland, Mingan Island and Gaspé Peninsula. 

The Preface, Table of Contents, How plants are used 
by the Nunavik Inuit, List of Vascular Plants, List of 


ENVIRONMENT 


Boreal Forest of Canada and Russia 
By W. O. Pruitt and L. M. Baskin. 2004. Pensoft Publishers, 
Sofia, Bulgaria.163 pages, Hardcover. $94.76 (Can.) 


Pruitt and Baskin are founders and directors of field 
research stations set up in the boreal forest regions of 
their respective countries. This book 1s far narrower in 
focus than the title suggests, however, concentrating on 
descriptions of the two field stations, their history, an 
outline of the work done there, and some observations 
on the surrounding terrain and landscape. The under- 
lying idea, though unstated except through the title, 
appears to be that these two small areas can be consid- 
ered representative of the vast regions of boreal forest 
on the two continents. The bulk of the book, though, 
does not deal with the forest but concentrates on its 
inhabitants. The text contains extensive discussion of 
the wildlife (pages 54-157) in the areas surrounding 
these field stations, with most of the discussion con- 
centrating on mammals, which are the principal research 
foci of both authors. It is written in a straight-forward 
narrative style, with few in-text citations, and is clearly 
aimed at the avocational naturalist or general reader. 

Most of the book focuses on the Taiga Biological 
Station, founded by Pruitt, which has been in opera- 
tion since 1973. The station is located northeast of 
Winnipeg, near the Manitoba-Ontario border. It has 
provided a base for many ecological and zoological 
studies through the years. The Station’s web page (www. 
wilds.mb.ca/taiga/intro.html) lists sixty-three written 
items, of which ten are MSc dissertations. The station 
has been used as a base for undergraduate field courses, 
visits by naturalists clubs, studies by museum curators, 
and projects by researchers from other institutions. Thus 
outreach and education, both formal and informal, are 
prime reasons for the station’s existence. 


Invascular Plants and General Index are all provided 
in three languages: French, English and Inuit. 

This book will be most interesting and useful to all 
visitors of the seventeen localities which were visited 
by the first author between 1983 and 1998 who made 
observations of the plants within a 10 km radius of 
each of the sites. It will also be very intriguing to any- 
one who has an opportunity to examine this beautiful 
book but never have an opportunity to visit the region. 


WILLIAM J. Copy 


Biodiversity, Program on Environmental Health, Agriculture 
and Agri-Food Canada, Wm. Saunders Building, Central 
Experimental Farm, Ottawa, Ontario, K1 A 0C6 Canada 


About a third of the text discusses the Kostroma 
Taiga Biological Station, located northeast of Moscow 
in the Volga River drainage basin, set up in 1977. From 
Baskin’s account, it appears that this area has been 
much more impacted by human activity than the area 
around the Canadian field station. He describes how 
intensive logging, game hunting, and fur trapping was 
carried out in the region. Recently, scientific activity 
at the field station has concentrated on game manage- 
ment and the study of animal populations under severe 
hunting pressure. Baskin also recounts attempts to in- 
troduce bison and cattle to the forest, to take advantage 
of grazing not being used by other large mammals 
(pages 97 -103). 

The majority of the text describes the wildlife of the 
two areas. Many of the animals (such as hares, wolves, 
bears, and moose) are common to both areas. Following 
a general survey of the wildlife and habitats, Baskin 
describes in more detail two species of birds and seven 
of mammals, mostly large mammals or fur-bearers. 
Pruitt contributes descriptions of thirteen mammals or 
groups of mammals, seven groups of birds, and a brief 
survey of amphibians, reptiles, and a few invertebrates. 
He devotes much attention to the ecology of Woodland 
Caribou (pages 109-116), which he describes “‘as the 
most endangered mammal in Canada today” (page 116), 
under pressure from fragmentation of and development 
in its habitat and with the potential for further shrink- 
ing of available habitat in response to climate change. 

Both field stations have been in operation for about 
a quarter century, yielding long-term data for their 
research areas. Such data sets are rare in ecology. Pruitt 
notes that he had the opportunity to set up studies of 
landscape recovery when the area around the station was 
afflicted by especially massive fires in 1980. Here, 


il 


however, is where I had a major problem with this 
book. There are no maps of the field sites! I found this 
omission truly surprising. It is impossible for anyone 
unfamiliar with the area to follow Pruitt’s discussion of 
the course of the fires through the region (pages 58-64). 
Similarly, Pruitt spends time describing six one acre 
study plots set up in the field area (pages 49-54). How- 
ever, I found myself wondering how these are distrib- 
uted on the landscape, how they relate to each other and 
other landscape features such as lakes and streams, and 
what the pattern of substrates might be. There is really 
not enough contextual information here to make this 
account intelligible. | had a similar problem with the 
discussion of the Russian field station. 

It is interesting to compare and contrast the difficul- 
ties both research directors describe in the running of 
their respective stations. The institutional and admin- 
istrative settings differ, but many of the problems are 
common to both. Pruitt deplores the depredations by 
forest companies and excoriates the provincial govern- 
ment for lack of support for conservation of the boreal 
landscape. He also makes an impassioned plea for the 
sustainable use of the boreal ecoregion, advancing “‘eco- 
tourism” as a use that is both sustainable and likely in 
the long-term to generate support for conservation poli- 
cies. Baskin laments the chaos following the breakup of 
the Soviet Union and describes the inroads unrestricted 
hunting, not for sport but for food and subsistence, made 
into the mammal populations around the field station. 
It is clear that both stations have struggled with uncer- 
tainty and under funding and have battled to keep going. 


THE CANADIAN FIELD-NATURALIST 


Vouk. TIS 


It is a tribute to the people involved that these stations 
have been operational for as long as they have. From 
Pruitt’s account, the spirit of camaraderie, companion- 
ship, and sense of community engendered in people who 
have worked at the field station is strong. Pruitt makes 
it clear that an attitude of “make do” rules at the Taiga 
Biological Station, with creative scrounging of materials 
and supplies from unlikely sources and a great deal of 
“sweat equity” contributed by students and volunteers. 
I expect that almost every researcher involved in a field- 
based science or natural history in Canada can only 
smile wryly in empathy. 

The most striking aspect of this book is the presen- 
tation of the information. The account is given in two 
languages with the text arranged in parallel columns, 
English to the left and Russian to the right, with equiv- 
alent paragraphs lined up. This is an interesting ap- 
proach, though I imagine that very few North American 
readers will be able to read both languages. Moreover, 
it makes the book twice as long as it needs to be and 
therefore probably increases its price. The book is 
well-produced, on high-quality glossy paper, with 78 
photographs, 72 of which are in colour, and robust bind- 
ing. However, the list price is outrageous for such a 
slender volume and, unfortunately, will probably severe- 
ly limit its distribution. 


ALWYNNE B. BEAUDOIN 


Royal Alberta Museum, 12845-102 Avenue, Edmonton Al- 
berta TSN OM6 Canada 


Democracy’s Dilemma: Environment, Social Equity, and the Global Economy 


By R Paehlke. 2004. MIT Press, Cambridge Massachusetts, 
306 pages. U.S.$ 16.95 Paper 


“Many have a sense that governments are increasing- 
ly out of control”. This book tackles this complex topic. 
It is written by one of the leading environmental thinkers 
and deals with major issues of globalization. Simply by 
assessing the global state of Forests, Topsoil, Biodiver- 
sity, Fisheries, Groundwater and Fossil Fuel, it becomes 
immediately obvious that we are borrowing the assets 
from future generations. As Paehlke states, these depres- 
sive facts are a logic conclusion when acting under the 
paradigm of “externalizing internal costs”, which is 
the result of a mis-lead economy promoted by the cur- 
rent view and definition of globalization. Here we get 
convincingly reminded about the significant political 
error made in the 90s that “unfettered” markets would 
be the only way to economic and all other forms of 
societal success. 

The book is divided into eight chapters and subsec- 
tions which convince that “the assumption that econom- 
ic expansion will fill our lives with sunshine” is faulty. 
As examples from India and U.S. indicate, wealthier 
nations are not the healthier ones. Nicely, the book des- 
cribes features of our current society: volunteer organ- 


izations and voting are in decline; political cynicism 
is all too normal due to the powerlessness of citizens. 

Besides a nice overview and introduction of global- 
ization issues, subsequent chapters deal with the media: 
the TV is at the heart of globalization, asking us per- 
manently to consume and to be entertained. Paehlke 
presents how this media is controlled by corporations, 
and thus not delivering conflicting or alternative mes- 
sages. Instead, it just focuses in a stereotypical way on 
natural disasters, accidents and arrests. Despite infor- 
mation technology, we are actually living in an age of 
“missing information”: Electronic media systematically 
avoids intellectual content in favor of visuals. Further, 
it is shown in this important book that the public Post 
Service deteriorated to an advertising bombardment 
infrastructure fueled by international corporations. In 
the U.S. alone, seventeen billion catalogues (64 for 
every man, woman and child) are mailed each year! 
Paehlke demands instead that the right of participatory 
access should be more often open to non-commercial 
interests and that individuals should have the right to 
establish some autonomy from commercial messages 
and images. 

One entire chapter deals with a “Three-Bottom-Line 
Perspective’ which eludes to the fact that “There has 


152 


not been any effective balancing of economic, social 
and environmental factors and interests”. For instance, 
GATT deals in its 20000 pages with corporate and 
business rights, but none deal with society and environ- 
mental issues. Other book sections elaborate on how to 
measure global progress: This matters as national GDPs 
go up, while there are strong indications that we are 
actually on the way backwards. For instance, tubercu- 
losis, a disease related to poverty and wiped out in ear- 
lier times, returned. In the year 2005 we are producing 
cloths to similar conditions as done in 1805. This books 
suggests alternatives to the narrow GDP metric such as 
an Ecosystem or Human Well-being Index; “Economic 
growth is a means, not the end”. 

Major social and democracy aspects are touched 
upon throughout this book. The author emphasizes 
that trade panels are empowered to overrule national 
environmental laws. Therefore, there is a strong need 
for them to be publicly accountable and elected. The 
reader will learn why the idea of a full-blown global 
government is flawed in principle: The challenge is to 
achieve global governance through the cooperation of 
effectively democratic national governments. Global- 
ization has flaws, for instance widely traveled products 
simply cannot be sustainable due to the high travel 
costs. We are exporting systematically environmental 
problems from rich to poor countries! Whereas U.S. has 
a high employment rate, in some countries it is even 
illegal to form unions. “In Mexico wages are a fraction 
of what they are in Canada and the U.S., effective pol- 
lution controls are more or less non-existent, and taxes 
for public health and education have been reduced or 
abolished”. Social costs of electronic capitalism are 
very real and not taken into account by those in busi- 
ness and government who make the relevant decisions. 

Paehlke actually shows that there are enough envi- 
ronmental and other warning signs that the current life 
style cannot be maintained, nor extended to third world 
countries. I liked best the book sections that deal with 
global economies and the environment, and with re- 
lated legal and policy issues. The CEC (Commission 
for Environmental Cooperation) in NAFTA is basically 
exposed as being powerless; CITES is widely ineffec- 
tive and has NOT stopped poaching. ITTO (Interna- 
tional Tropical Timber Organization) as well as the Bio- 
diversity Convention have basically failed by not halt- 
ing tropical forest loss. RAMSAR and the Bonn Con- 
vention on Migratory Species are unfortunately not 
referred to in this book but are known to be inefficient 
as well. International environmental laws suffer from 
enforcement and binding rules, whereas the economic 
counterparts always offer drastic punishments when any 
of their terms are violated. 

NAFTA, WTO and GATT have policies that allow 
polluters to sue national and local governments for even 
attempting to protect the environment. IMF (Interna- 
tional Montary Fund) deals only with narrow domestic 
economic interests. Paehlke gives us nice examples such 
as the large pier in Cozumel threatening the Paradiso 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Reef (CEC jurisdiction), or the infamous dolphin-tuna 
ruling where the U.S. Marine Mammal Protection Act 
violated GATT in Mexico and was ruled not to be valid 
outside of national jurisdiction. Another case is men- 
tioned where the WTO is in conflict with sea turtles 
and instead puts trade first. At least the ISO has some 
environmental management standards, but then this 
organization is again administrated by the corporate 
industry themselves. Why are environmentalists not 
asked when it comes to taxation or annual budgets? Per- 
haps we need a tax on consumption and energy waste? 

The Ironies of the Global Age are: Rising poverty in 
the face of an enormous surge in productive capacities, 
declining leisure time in the face of increasingly auto- 
mated industrial production, and reversals in environ- 
mental protection in the face of advancing environ- 
mental knowledge and high levels of environmental 
concern. This book fully exposes this “Cult of Impo- 
tence”, which now is so widely found in governments 
and responsible agencies in the world. Critical voices 
are not wanted, discriminated against. It is indeed true, 
as stated by the author, that major administrative con- 
cepts such as communism, failed due to ignoring envi- 
ronmental issues and by being undemocratic! 

The reader gets presented with a balanced view 
though. For instance that Globalization has seen mod- 
est successes in regards to human rights, labor rights 
and the rights of women and children. Computer and 
WWW spread democracy. Thirdly, the California effect 
(an increase in economy results into an improved 
environmental standards; e.g., air) is shown. 

This book deals with the substance of democracy, 
not with its hollow form. The 275 pages are full of 
fascinating bits and pieces: What’s the meaning of the 
(working) life? We replaced tedious industrial work 
with tedious and pointless marketing and retail work 
spending a considerable percentage of today’s employ- 
ees who are devoted to the task of selling us things we 
do not otherwise even imagine we need. Exotic species 
problems as a direct function of Globalization. Of 
interest is also the statistic that during the last 50 years 
humans have used more resources than during the 
entire previous human civilization. It is fascinating to 
learn that one can hire 47 Philipinos for 1 French 
worker, or that 200 of the wealthiest individuals are as 
rich as 41% of the world population! Some readers might 
be surprised to learn that U.S. has replaced Japan when 
it comes to workload. Subsequently, divorce rates and 
family breakdowns are on a record high. I am really 
unclear why such a hard working nation has the high- 
est energy consumption though. 

Besides fascinating facts, I also like the conclusions 
brought forward by Paehlke: Other authors described 
Globalization already as the “environmental race to the 
bottom”. It is referred to as “laissez faire” capitalism. 
History shows that “laissez faire” cannot easily be re- 
formed: It needed the disasters of the Great Depression 
and of World War 2 to shake the hold of an earlier ver- 
sion of the free market orthodoxies on western govern- 


155 


ments and societies. Nowadays, Global Change and 
Poverty are probably the single biggest failures of the 
free market system. 

In this book, I found no major shortcoming (other 
than that Germany does not have a 48h working week, 
modern Russia and China get hardly mentioned, and 
the book index is incomplete), but some of the thoughts 
presented by the author could be challenged. It is cor- 
rect that all natural commodities (mining, forestry, 
agriculture and fish products) but oil are going down 
in value. However, likely these commodities just loose 
due to the dominance, and convenience brought by, 
oil. Unfortunately, despite its catching and fascinating 
subject, this text makes for a hard and long read: I find 
it unnecessarily boring and repetitive. 

In conclusion, we lack a global citizen movement. 
The reader of this book will whole heartedly agree that 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


globalization also requires the definition of minimum 
standards for welfare, environment, taxation and wages. 
We are left with the need for crucial reforms: (1) we need 
a media reform, (ii) a social science reform away from 
“economics only”, (iii) a radical election campaign 
finance reform, (iv) progress in the reduction of total 
work time with families at least to the level that was 
normal prior to the decline in civic participation, (v) 
and an appreciation of the need for global governance 
rooted in restored democracy at the level of the 
nation-state. | recommend reading this book and get- 
ting into action. 


FALK HUETTMANN 


Institute of Arctic Biology, Biology & Wildlife Department, 
University of Alaska-Fairbanks, Alaska 99775 USA 


Emulating Natural Forest Landscape Disturbances: Concepts and Applications 


Edited by Ajith H. Perera, Lisa J. Buse, and Michael G. Weber. 
2004. Columbia University Press, New York. 315 pages. 


Disturbance is ubiquitous in forest ecosystems. Dis- 
turbed by the extremes of either catastrophic, stand- 
replacing events including fire, insect herbivory and 
extensive wind throw, or periodic, small-scale gap 
processes mediated by fungal pathogens, forests are 
in constant flux when viewed from a long-term, land- 
scape perspective. A wide range in the periodicity, in- 
tensity and scale of disturbance events, and in the diver- 
sity of bio-edaphic interactions creates a complex, fluid, 
heterogeneous forest landscape. 

Practitioners of sustainable forest management have 
accepted the essential links among natural disturbances, 
forest and stand structural heterogeneity and organism 
biodiversity. Recognition of these links has generated 
the conceptual and empirical development of a natural 
disturbance-based forest management defined as “an 
approach in which forest managers develop and apply 
specific management strategies and practices, at ap- 
propriate spatial and temporal scales, with the goal of 
producing forest ecosystems as structurally and func- 
tionally similar as possible to the ecosystems that would 
result from natural disturbances, and that incorporate 
the spatial, temporal, and random variability intrinsic 
to natural systems.” (page 4) 

Widely accepted is the assumption 1s that forest bio- 
logical, structural and functional diversity developed 
within the boundaries defined by natural disturbances. 
As such, human interventions, such as logging or the 
use of prescribed fire, that conform “more or less” to the 
temporal and spatial dimensions of natural, historic dis- 
turbances are deemed to be those most successful in 
conserving biological diversity. 

The book’s editors include a forest landscape ecol- 
ogist (Perera) and a forest biologist (Buse), both from 
the Ontario Forest Research Institute, and a fire ecol- 
ogist (Weber) from the Great Lakes Forestry Centre of 


the Canadian Forest Service. Together they have pro- 
duced a significant volume with contributions from 
both practitioners and academics who are actively en- 
gaged in the development of natural disturbance-based 
forestry. 

The collection of essays is broken down into three 
main sections. The first section deals with the theoret- 
ical and conceptual foundations of emulating natural 
disturbance in forest management. The authors of the 
five essays in this first section do a commendable job 
of describing the ecological foundations and biodiver- 
sity implications of emulating natural disturbance in 
forest management. The eight contributors to the second 
section treat a host of case studies from different forest 
ecosystems throughout the United States and Canada. 
With the help of computer simulation models, historic 
natural disturbance regimes are assessed for use in dis- 
turbance-based forest management. These empirically- 
based technical studies are site- and computer-model 
specific and provide state of the art concrete applications 
of natural disturbance-based forestry. A 16-page suite 
of computer-generated full-colour maps provides exam- 
ples of output from the various simulation models. The 
final section composed of seven chapters addresses the 
actual feasibility and practice of emulating natural dis- 
turbance through forest stand-based management. In- 
dustry and environmental non-governmental perspec- 
tives are also treated in this final section. The final 
chapter is an excellent synthesis of the current state of 
knowledge about emulating natural disturbance in north- 
ern North American forests. 

Palaeoecological and historical evidence reveal that 
many forest ecosystems are shaped by the interaction 
of climate change, natural disturbances and human activ- 
ities. The emulation of natural disturbance, therefore, 
can be akin to shooting at a constantly moving target. 
This book does not present emulation of natural dis- 
turbance as a forest management panacea. Nor does one 


154 


get a sense that forest managers must slavishly follow 
natural patterns and processes. Emulation of natural 
disturbance provides a sensible guide to sustainable 
forest management. It cannot, however, be carved in 
stone. The dynamic nature of forests, the existence of 
multiple successional pathways in response to distur- 
bance and the even more dynamic needs and values of 
human societies would prevent that. The last lines of the 
book say as much: “Over the long term, the ultimate 
success of the approach will be determined by the 
answers to the questions of whether it 1s ecologically 
superior to other forest management paradigms, eco- 
nomically feasible for forestry practitioners, and social- 
ly acceptable.” (page 274) 

The editors consider the book well suited to all forestry 
professionals including practitioners, policy makers 


MISCELLANEOUS 


The Nature Journal: A Handbook 


By Frederick W. Schueler and Aleta Karstad. 2004. Bishops 
Mills Natural History Centre, RR 2, Oxford Station, Ontario 
KOG ITO and Little Ray’s Reptile Zoo, 5305 Bank Street, 
Ottawa Ontario KIX 1H2 [www.thenaturejournal.ca]. 
$40.00 


For any naturalist, whether professional or hobbyist, 
some form of notes are an essential reference to past 
observations. At one level they can serve for compar- 
ison with new observations or as reminders of where 
and when to look for certain species or seasonal phe- 
nomena. At another, they can be the basis for prepara- 
tion of accurate published accounts. How to format and 
organize them for permanence and accessibility has 
more solutions than observers as even individual sys- 
tems evolve over time. For those just starting to keep 
records or dissatisfied with previous efforts, Fred and 
Aleta Schueler offer a solution. 

Since the 1960s, one or both have been observers and 
commentators of nature, collectors of biological spec- 
imens for museums, and writers and illustrators of both 
popular accounts and scientific papers. They have re- 
corded their raw field data in a variety of journal and 
data formats. Here, they attempt to standardize this 
experience and outline a universal system that will 
produce notes not only of use to observers themselves 
but also serve as a permanent record of value for others. 
The Schueler’s rightly regard leaving a useable record 
for future naturalists as a prime responsibility of us all. 

What is outlined here is partly adapted from the 
once widely used system credited to Joseph Grinnell, 
Curator of the Museum of Vertebrate Zoology at 
Berkeley, California, in the first half of the 20" centu- 
ry. It also draws on field data entry sheets developed 


THE CANADIAN FIELD-NATURALIST 


Vol. Tf 


and researchers. To this list I would add conservation 
biologists, environmentalists and even environmental 
philosophers. The book deals not only with the con- 
ceptual and practical considerations of the emulation 
of natural disturbance, but helps to raise broader ques- 
tions about biodiversity conservation in dynamic forests 
subject to both natural and human-induced change. 
This book should stimulate those philosophical ques- 
tions, but you will have to resort to other works for the 
beginnings of a response. 


JOHN MCCARTHY 


Ecology Project, Jesuit Centre for Social Faith and Justice, 
P.O. Box 1238, Guelph, Ontario N1H 6N6 Canada 


at the National Museum of Canada (now the Canadian 
Museum of Nature) in ichthyology and herpetology in 
the 1960s and 1970s. 

A sturdy seven-ring binder is prefixed with 37 pages 
of instruction including: The role of archival natural- 
ists, What needs study? What should I look for? What 
should I record? What if I do not know its name? Who 
is interested in what I observe? and explanation: The 
pages, Archival materials, Fine tuning accounts, Inter- 
preting the datasheet, The Grinnell System: a brief 
history, Notes on journal-keeping, Acknowledgments, 
and Resources. An irritation 1s that an included reprint- 
ing of an unpublished report with three references has 
no documentation for these. 

The Mamre Journa/ contains acid-free archival paper 
of 50% cotton (Ernscliffe Linen Bond): 30 lined journal 
pages, 15 pocket pages, 15 “catalog” pages, 15 species 
account pages, 15 datasheets, 30 blank pages for draw- 
ings, 2 heavy acid-free pages for watercolour or labels, 
4 acid-free separators. Also included is one archival ink 
“Pigma” felt tip pen and a ruled plastic page finder. 
An enthusiastic field-naturalist would soon use up the 
initial stock but additional pages are available from 
the authors. 

In promised progress is an electronic data base for 
field notes. Although, as is pointed out here, this 1s far 
easier to search later but can be more time-consuming 
than written notes. The web address www.thenature- 
journal.ca will provide updates. 


FRANCIS R. COOK 


Researcher Emeritus, Canadian Museum of Nature, Ottawa, 
Ontario K1P 6P4 Canada 


i> 


NEw TITLES 


+Available for review * 
Assigned for review 


Zoology 


Evolution of Amphibian Skull. Advances in Amphibian 
Research, Volume 9. By N. Lebedkina. 2005. Pensoft Pub- 
lishers, Sofia 1113, BULGARIA, Acad. G. Bonchev str. bl.6. 
In English, 265 pages. URO 44.50 Paper. (Surface mail deliv- 
ery: URO 5. Airmail delivery overseas: URO 15.) 


*The Birds of Azerbaijan. M. Patrikeev, 2004. Pensoft 
Publishers, Sofia-Moscow. 380 pages. URO 68.80 — Surface 
mail delivery: URO 5. Airmail delivery overseas: URO 19. 


A Field Guide to the Caterpillars. By T. Allen, J. Brock, and 
J. Glassberg. 2005. Oxford University Press, 70 Wynford Drive, 
Don Mills, Ontario, M3C 1J9. 224 pages. Can $34.95. 


Drawing Birds. By J. Busby. 2005. Timber Press, 133 SW 
2™ Avenue, Suite 450, Portland, Oregon, USA. 144 pages, 
Paperback, U.S.$19.95. 


Hunting for Frogs on Elston, and Other Tales from Field & 
Street. By Jerry Sullivan, Edited by Victor M. Cassidy. 2004. 
University of Chicago Press, 5801 Ellis Avenue, Chicago, 
Illinois 60637 USA. 320 pages. U.S.$25.00 cloth. 


The Origin and Evolution of Mammals. By T. S. Kemp. 
2004. Oxford University Press, 70 Wynford Drive, Don Mills, 
Ontario, M3C 1J9. 342 pages. £80.00 cloth. 


Noncooperative Breeding in the California Scrub Jay. By 
W. Carmen. 2004. Cooper Ornithological Society. 100 pages. 
U.S.$15 Paper. 


Pronghorn: Ecology and Management. By B. O’ Gara and 
J. Yoakum. 2004. University Press of Colorado, 5589 Arapahoe 
Road, Suite 206C, Boulder, Colorado 80303 USA. 904 pages. 
U.S.$85. Cloth. 


Shorebirds of North America — the photographic guide. 
By Dennis Paulson. 2005. Princeton University Press, 41 
William Street, Princeton, New Jersey 08540 USA. 384 pages. 
U.S.$29.95 paper, $65 cloth. 


*Waterfowl of Eastern North America. By Chris Earley. 
2005. Firefly Books, 66 Leek Crescent, Richmond Hill, On- 
tario, L4B 1H1. 158 pages. $19.95 paper $29.95 Cloth. 


White as a Ghost — Winter Ticks and Moose. By Bill Samuel. 
2004. Federation of Alberta Naturalists, 11759 Groat Rd., 
Edmonton, Alberta TSM 3K8. 99 pages. Can. $24.95. 


Wheatears of the Palearctic. Ecology, Behaviour and Evo- 
lution of the Genus Oenanthe. By E. Panov, 2005. Pensoft 
Publishers, Sofia-Moscow, 439 pages. Cloth URO 58.80 — 
Surface mail delivery URO 7. Airmail delivery overseas 
URO 14. 


THE CANADIAN FIELD-NATURALIST 


Volum 


Botany 
California’s Wild Gardens — A Guide to Favorite Botanical 
Sites. Edited by Phyllis M. Faber University of California 
Press, 2120 Berkeley Way, Berkeley, California 94704-1012 
USA. 248 pages. U.S.$34.95. 


*An Illustrated Guide to the Eastern Woodland Wild- 
flowers and Trees. By M. Choukas-Bradley. 2004. University 
of Virginia Press, P.O. Box 400318, Charlottesville, Virginia 
22904-4318 USA. 424 pages. U.S.$39.85. Cloth. 


Other 


Curious by Nature. By Candace Savage. May 2005. Grey- 
stone Books, #201 — 2323 Quebec Street, Vancouver, British 
Columbia, V5T 4S7. 160 pages, $22.95 Can. Paper. 


+ The Charting of an Atmospheric Environment. By M. 
Hamilton. 2004. Vantage Press Inc., 419 Park Avenue South, 
New York, New York 10016 USA. 262 pages, U.S.$24.95. 


*The Earth’s Blanket — Traditional Teachings for Sus- 
tainable Living. By N. Turner. 2005. Douglas & McIntyre 
Publishing Group Suite 500, 720 Bathurst Street, Toronto, 
Ontario MSS 2R4. 304 pages, U.S.$29.95 Cloth. 


Ecological Methods in Forest Pest Management. By David 
Wainhouse. 2004. Oxford University Press, 70 Wynford Drive, 
Don Mills, Ontario M3C 1J9. 248 pages, £55.00 Cloth. 


Geoffroy Saint-Hilaire: A Visionary Naturalist. By Herve 
Le Guyader, Translated by Marjorie Grene. 2003 University 
of Chicago Press, 801 Ellis Avenue, Chicago, Illinois, 60637 
USA. 288 pages, U.S. $45.00 Cloth. 


Atlas of Pacific Salmon — The First Map-Based Status 
Assessment of Salmon in the North Pacific. By Xanthippe 
Augerot. 2005. University of California Press, 2120 Berkeley 
Way, Berkeley, California 94704-1012 USA. 161 pages, 
U.S.$34.95. 


Preserving the Living Past — John C. Merriam’s Legacy 
in the State and National Parks. By Stephen R. Mark 
University of California Press, 2120 Berkeley Way, Berkeley, 
California 94704-1012 USA. 219 pages. U.S.$39.95. 


Record of Documents, 1964 — 2004, Pertaining to the Pre- 
servation of Kamoka Park, ON with a biography of Os- 
mund Langvet (1926 — 1979). By W. Judd. Phelps Publish- 
ing Company London, Ontario. 


Catalogue of Meetings 1988-2004 of the Birding Wing of 
the Mcllwraith Field Naturalists of London ON. By W. 
Judd. Phelps Publishing Company London, Ontario. 


Children’s Books 


The Kids’ Guide to Zoo Animals. By Michelle Gilders. 2004. 
Red Deer Press, Trailer C, 2500 University Drive, Calgary 
Alberta, T2N 1N4 Canada. 264 pages, cloth, $16.95 Can. 


News and Comment 


Marine Turtle Newsletter (107) 


January 2005. 28 pages: ARTICLES: Morphometric 
Analysis of the Northern Subpopulation of Cavefa caretta 
in South Carolina, USA — Marine Turtle Conservation in 
Viet Nam — Towards 2010 — Notes: Holding a Live 
Leatherback Turtle in Israel: Lessons Learned — The 
Marine Turtles of Hinatuan Bay, Surigao del Sur, Mindanao, 
Philippines — Two Records of Live Olive Ridleys from 
Central California, USA — Oceanic Habitats for Loggerhead 
Turtles in the Mediterranean Sea — Sea Turtle Nesting Sites 
at Lian, Batangas, Philippines — New Records on the Dis- 
tribution of Loggerhead Turtles (Caretta caretta) in the 
Philippines — Stranded Sea Turtles on the Coast of Paraiba 
— Brazil — Underwater Oviposition by a Hawksbill Turtle 


in Guadeloupe, French West Indies — IUCNMTSG UppaTEe 
— ANNOUNCEMENTS — NEWS & LEGAL BRIEFS — RECENT 
PUBLICATIONS. 

The Marine Turtle Newsletter is edited by Brendan J. 
Godley and Annette C. Broderick, Marine Turtle Research 
Group, Centre for Ecology and Conservation, University of 
Exeter in Cornwall, Tremough Campus, Penryn TRIO 9EZ 
United Kingdom; e-mail MTN @seaturtle.org; Fax +44 1392 
263700. Subscriptions and donations towards the production 
of the MTN can be made online at <http://www.seaturtle.org/ 
mtn/> or postal mail to Michael Coyne (online Editor) Marine 
Turtle Newsletter, 1 Southampton Place, Durham, North 
Carolina 27705 USA (e-mail: mcoyne @seaturtle.org). 


se eal Newsletter of the Declining Amphibian Populations Task Force (67): Last Printed 
e 


Number 67 February 2005. *Last Printed Issue* Contents: 
DAPTE Seed Grants 2005 — Action for the axolotl at Lake 
Xochimilco [Richard A. Griffiths and Ian G. Bride] — The 
Stutus of Great Crested Newt Breeding Sits in Serbia [Jelka 
Crnobrnja-Isailovic, Ivan Aleksic & Jan Willem Arntzen, 
DAPTFE Seed Grant Recipients — Meeting on disease and 
threats to amphibian biodiversity [James Collins] — Introduced 
fish, rotan Perco//lus glenii — an unavoidable threat for Euro- 
pean amphibians [Andrey N. Reshetnikov] — Froglog Shorts: 
Due to escalating printing and shipping costs the printed 
copies of Froglog will no longer be be generally distributed. 
The newsletter /7og/og will continue, posssibly increasing in 
size and frequency, available free at http://www.open.ac.uk/ 
daptf/froglog. It can also be accessed through Herp Digest’s 
listserve at http://www.herpdigest.org/ 


New Canadian Journal: Wildiife Afield (\ and 2) 


A new British Columbia semi-annual publication Wi/d/ife 
Afteld ISSN 1712-2880) is to be issued by The Biodiversity 
Centre for Wildlife Studies twice a year. 

Volume 1, Number 1, contains FROM THE EpiTor: Wi/d/ife 
Afie/d — An introduction; On the covers — FEATURE ARTICLE: 
Food habits of the Barn Ow] in the southern interior of British 
Columbia (Linda M. Van Damme and Mark Nyhof — NoTEs: 
First occurrence of Wandering Salamander on the Sunshine 
Coast of British Columbia (Glenn R. Ryder and R. Wayne 
Campbell) — Dusky Flycatcher breeding in the Peace River 
Region of British Columbia (Doug Brown) — Upland Sand- 
piper breeding near Chetwynd, British Columbia (R. Wayne 
Campbell) — Occurrence of the Rock Wren on Vancouver 
Island and other islands in the Strait of Georgia (Michael I. 
Preston) — Unsuccessful introduction of the California Quail 
to the Queen Charlotte Islands (R. Wayne Campbell) — wILp- 
LIFE DATA CENTRE: Featured species — Semiplamated Plover 
(R. Wayne Campbell) — Report of the Wildlife Data Centre: 
1 June 2003 to 30 June 2004 (Michael I. Preston) — BRITISH 
COLUMBIA ROUND-UP: Field notes and caring people — News 


Frog/og is the bi-monthly newsletter of the Declining Am- 
phibian Populations Task Force of The World Conservation 
Union (IUCN)/Species Survival Commission (SSC) and is 
supported by The Open University, The World Congress of 
Herpetology, and Arizona State University. It is edited by 
Jeanne McKay (65) and Jeanne McKay (66),, Department of 
Biological Sciences, The Open University, Walton Hall, Milton 
Keynes, MK7 6AA, United Kingdom; e-mail: daptf@ open. 
ac.uk. Funding for /vog/og is underwritten by the Detroit 
Zoological Institute, P. O. Box 39, Royal Oak, Michigan 
48068-0039, USA. Publication is also supported by Peace 
Frogs www.peacefrogs.com and by RANA and the US 
National Science Foundation grants DEB-0130273. 


of Friends — From the Archives — Final Flight — Publica- 
tions of Interest — Announcements and Meetings — Support 
for a Shared Vision — Your Data at Work — Biodiverstiy 
Centre for Wildlife Sudies: Our Logo. 

Volume 1, Number 2, contains: FROM THE Epiror: And the 
ship sets sail — On the covers — FEATURE ARTICLES: Temporary 
colonization of Cleland Island, British Columbia, by Common 
Murres from 1969-82 (Harry R. Carter) — Conservation pri- 
orities and peripheral species in the south Okanagan: Con- 
siderations for a proposed National Park (Joanna Preston) 
— Effectiveness of global protected areas? Perspectives for 
British Columbia (Michael I. Preston) — Nores: Observations 
of breeding Dusky Flycatchers in the central Okanagan Valley, 
British Columbia (Chris Charlesworth) — Field observations 
of Bullfrog (Rana catesbeina) prey in British Columbia (R. 
Wayne Campbell and Glenn R. Ryder) — Gross bill de- 
formity and longevity in a Northern Flicker (Sherry L. Lid- 
stone) — Gray Wagtail (oracilla cinerea): a new species 
for British Columbia (Jerry Etzkorn and Janet Etzkorn) — 
Lark Sparrow nesting in the Peace River region of British 


156 


2004 


Columbia (Steve Myers) — Gray Wolf predation on Trumpeter 
Swan eggs (R. Wayne Campbell) — Weather influences 
parenting among Red-necked Grebes and Western Grebes 
on Duck Lake, Creston Valley (Linda M. Van Damme) — 
WILDLIFE DATA CENTRE: Featured species — Wood Frog (R. 
Wayne Campbell) — Report of the Wildlife Data Centre: 1 
July 2004 to 31 December 2004 (Michael J. Preston) — 
BRITISH COLUMBIA ROUND-UP: Field notes for caring people 
— News of Friends — From the archives — Publications of 


NEws AND COMMENT 


137 


interest [Compiled and edited by Chris Siddle] — Announce- 
ments and meetings. 

Annual memberships in the Biodiversity Centre for Wildlife 
Studies (which includes Wi/d/ife Afie/d) are $30 (individual), 
$40 (family). $20 (student), $500 (life). Biodiversity Centre 
for Wildlife Studies, PO Box 6218, Station C, Victoria, 
British Columbia V9P 5L5 Canada. Tel/Fax: 250-477-0465; 
e-mail: editor @ wildlifebc.org. 


158 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Advice for Contributors to 7e Canadian Field-Naturalist 


Content 

The Canadian Field-Naturalist is a medium for the publi- 
cation of scientific papers by amateur and professional natu- 
ralists or field-biologists reporting observations and results 
of investigations in any field of natural history provided that 
they are original, significant, and relevant to Canada. All read- 
ers and other potential contributors are invited to submit for 
consideration their manuscripts meeting these criteria. The 
journal also publishes natural history news and comment items 
if judged by the Editor to be of interest to readers and sub- 
scribers, and book reviews. Please correspond with the Book 
Review Editor concerning suitability of manuscripts for this 
section. For further information consult: A Publication Policy 
for the Ottawa Field-Naturalists’ Club, 1983. 7ie Canadian 
Field-Naturalist 97(2): 231-234. Potential contributors who 
are neither members of 7he Ottawa Field-Naturalists’ Club 
nor subscribers to 7e Canadian Field-Naturalist are encour- 
aged to support the journal by becoming either members or 
subscribers. 


Manuscripts 

Please submit, to the Editor, in either English or French, 
three complete manuscripts written in the journal style. 
The research reported should be original. It is recommended 
that authors ask qualified persons to appraise the paper before 
it is submitted. All authors should have read and approved it. 
Institutional or contract approval for the publication of the data 
must have been obtained by the authors. Also authors are ex- 
pected to have complied with all pertinent legislation regard- 
ing the study, disturbance, or collection of animals, plants or 
minerals. The place where voucher specimens have been de- 
posited, and their catalogue numbers, should be given. Lati- 
tude and longitude should be included for all individual local- 
ities where collections or observations have been made. 

Print the manuscript on standard-size paper, doublespace 
throughout, leave generous margins to allow for copy mark- 
ing, and number each page. For Articles and Notes provide 
a citation strip, an abstract and a list of key words. Gener- 
ally, words should not be abbreviated but use SI symbols for 
units of measure. The names of authors of scientific names may 
be omitted except in taxonomic manuscripts or other papers 
involving nomenclatural problems. “Standard” common names 
(with initial letters capitalized) should be used at least once 
for all species of higher animals and plants; all should also 
be identified by scientific name. 

The names of journals in the Literature Cited should be 
written out in full. Unpublished reports and web documents 
should not be cited here but placed in the text or in a sepa- 
rate Documents Cited section. List the captions for figures 
numbered in arabic numerals and typed together on a separate 
page. Present the tables each titled, numbered consecutively 
in arabic numerals, and placed on a separate page. Mark in 
the margin of the text the places for the figures and tables. 


Check recent issues (particularly Literature Cited) for 
journal format. Either “British” or “American” spellings are 
acceptable in English but should be consistent within one 
manuscript. The Oxford English Dictionary, Webster’s 
New International Dictionary and le Grand Larousse 
Encyclopédique are the authorities for spelling. 


Illustrations 

Photographs should have a glossy finish and show sharp 
contrasts. Electronic versions should be high resolution. Photo- 
graphic reproduction of line drawings, no larger than a 
standard page, are preferable to large originals. Prepare line 
drawings with India ink on good quality paper and letter (don’t 
type) descriptive matter. Write author’s name, title of paper, and 
figure number on the lower left corner or on the back of each 
illustration. 


Reviewing Policy 

Manuscripts submitted to 7he Canadian Field-Naturalist 
are normally sent for evaluation to an Associate Editor (who 
reviews it or asks another qualified person to do so), and at 
least one other reviewer, who is a specialist in the field, cho- 
sen by the Editor. Authors are encouraged to suggest names 
of suitable referees. Reviewers are asked to give a general 
appraisal of the manuscript followed by specific comments 
and constructive recommendations. Almost all manuscripts 
accepted for publication have undergone revision—sometimes 
extensive revision and reappraisal. The Editor makes the 
final decision on whether a manuscript is acceptable for pub- 
lication, and in so doing aims to maintain the scientific quality, 
content, overall high standards and consistency of style, of 
the joumal. 


Special Charges — Please take note 

Authors must share in the cost of publication by paying 
$80 for each page, plus $15 for each illustration (any size up 
to a full page), and up to $80 per page for tables (depending 
on size). Authors may also be charged for their changes in 
proofs. Reproduction of color photos is extremely expensive; 
price quotations may be obtained from the Business Manager. 
If grant or institutional funds are not available, club members 
and subscribers may apply for a waiver of charges for the 
first five pages. 

Limited joumal funds are available to help offset publi- 
cation charges to authors with minimal financial resources. 
Requests for financial assistance should be made to the Busi- 
ness Manager when the manuscript is accepted. 


Reprints 
An order form for the purchase of repents will accompany 
the galley proofs sent to the authors. 


FRANCIS R. Cook, Editor 
RR 3 North Augusta, Ontario KOG IRO Canada 


TABLE OF CONTENTS (concluded) Volume 119 Number 1 2005 
Feature Article 
Why and how to study a snowcover WILLIAM O. PRUITT, JR. 118 
Notes 
Découverte de la salamandre a quatre doigts, Hesidactylium scutatum, a Québec, Québec: 
limite nord-est de l’espéce sur la rive nord du fleuve Saint Laurent 
DANIEL POULIOT et JEAN-FRANCOIS DESROCHES 129 
Use of a bridge for day roosting by the Hoary Bat, Zasturus cinereus 
PAUL HENDRICKS, JOSEPH JOHNSON, SUSAN LENARD, and COBURN CURRIER 132 
Aggressive behaviour exhibited by a San Joaquin Swift Fox, Va/pes macrotis mutica 
HOWARD O. CLARK, JR. 134 
n ancient Wolf, Carus /upus, den and associated human activity in the southwestern Yukon Territory 
RICHARD FARNELL, P. GREGORY HARE, and DANIEL R. DRUMMOND Me 
apillate Watermeal, Wo/ffia brailiensis in eastern Ontario: An addition to the flora of Canada 
E. R. THOMSON [37 
ecord size female Coyote, Canis /atrans JONATHAN G. Way and ROBERT L. PROIETTO 139 
k Reviews 
OOLOGY: Amphibiens et reptiles du Quebec et les maritimes — Mammals of Australia — Birds of the 
Raincoast: Habits and Habitat — The Bird Almanac: A Guide to Essential Facts and Figures of the 
World’s Birds — Common Birds of Ontario — Frogs of Australia: An Introduction to their 
Classification, Biology and Distribution — Experimental Approaches to Conservation Biology — 
Gulls of North America, Europe, and Asia — The Monarch Butterfly: Biology and Conservation — 
The Behavior and Ecology of Pacific Salmon and Trout — Venomous Reptiles of the Western Hemisphere 
— Whales and Dolphins of the World 141 
OTANY: Atlas des Plantes Villages du Nanavik — Atlas of Plants of the Nunavik Villages 150 
NVIRONMENT: Boreal Forest of Canada and Russia — Democracy’s Dilemma: Environment, Social 
Equity, and the Global Econony — Emulating Natural Forest Landscape Disturbances: Concepts 
and Applications 150 
ISCELLANEOUS: The Nature Journal: A Handbook 154 
EW TITLES 155 
ews and Comment 
arine Turtle Newsletter (107) — Froglog (67) February 2005: last printed issue — New 
Canadian Journal: Wi/dlife Afield(\ & 2) 156 
Advice to Contributors 158 


Mailing date of the previous issue 118(4): 3 March 2006 


THE CANADIAN FIELD-NATURALIST Volume 119 Number 1 


Articles 


Cronology of range expansion of the Coyote, Canzs /atrans, in New York 
HEATHER M. FENER, JOSHUA R. GINSBERG, ERIC W. SANDERSON, and MATTHEW E. GOMPPER 


Nesting behavior, ecology, seasonal and geographic variation of the Sand Wasp, 
Stucttella emarginata (Hymenoptera, Sphecidae) FRANK E. KURZEWSKI and HUGH F. BOYLE 


Environment and distribution of age 0 fishes in River Canard, a lowland Ontario river 
JOHN K. LESLIE and CHARLES A. TIMMINS 


Charophytes of insular Newfoundland II: Chara evoluta and Chara canescens 
HENRY MANN and E. M. V. NAMBUDIRI 


The effect of human activity on ant species (Hymenoptera: Formicidae) richness 
at the Mont St. Hilaire Biosphere Reserve, Quebec 
JONATHAN Z. SHIK, ANDRE FRANCOEUR, and CHRISTOPHER M. BUDDLE 


Seasonal diets of Newfoundland Martens, Wartes americana atrata 
JOHN W. GOOSE and BRIAN J. HEARN 


Pollination and breeding system of Lowbush Blueberries, Vaccinium angustifolium Ait. 
and V. myrtilloides Michx. (Ericacaeae) in the boreal forest 
MASAYUKI UsulI, PETER G. KEVAN, and MARTYN OBBARD 


Productivity of Osprey, Pandion haliaefus, nesting on natural and artificial structures 
in the Kawartha Lakes, Ontario, 1991-2001 
PAMELA A. MARTIN, SHANE R. DE SOLLA, PETER J. EWINS, and MICHAEL E. BARKER 


Effects of wetland creation on breeding season bird use in boreal eastern Ontario 
Davip A. Locky, J. CHRIS DAVIES, and BARRY G. WARNER 


Lichen trimlines in Peace-Athabasca Delta: Variations in flora, form, and disturbance regime 
KEVIN P. TIMONEY and JANET MARSH 


Identification of a marine green alga Fercursaria percursa from hypersaline springs 
in the middle of the North American continent 
KATHLEEN L. LONDRY, PASCAL H. BADIOU, and STEPHEN E. GRASBY 


New distributional records of Ca//igrapha species (Leaf Beetles) in North America 
(Coleoptera, Chrysomelidae, Chrysomelinae) JESUS GOMEZ-ZURITA 


Weight-carrying ability and caching behavior of Gray Jays, Pertsoreus cacadensis. 
Adaptions to boreal winter LYNN L. ROGERS 


Premieres mentions et réparation de la Salamandre sombre du Nord, Desmognathus Juscus, 
sur la rive nord du fleuve Saint-Laurent, au Québec 
JEAN-FRANCOIS DESROCHES et DANIEL POULIOT 


First records of Long-beaked Common Dolphins, Dedphinus capensis, in Canadian waters 
JOHN K. B. FORD 


Occurrence, composition and formation of Ruppia, Widgeon Grass, 
balls in Saskatchewn Lakes 
RANDY W. OLSON, JOSEPH K. SCHMUTZ, and U. THEODORE HAMMER 


ISSN 0008-3550 


(continued on inside back cove 


The CANADIAN 
FIELD-NATURALIST 


Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada 


IBRAR* 


Volume 119, Number 2 April—June 2005 


The Ottawa Field-Naturalists’ Club 


FOUNDED IN 1879 


Patrons 
Her Excellency The Right Honourable Adrienne Clarkson, C.C., C.M.M., C.D. 
Governor General of Canada 
His Excellency John Ralston Saul, C.C. 


The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to _ 
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields 
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ- 
ments of high quality for living things. 


Honorary Members 


Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope 
Charley D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr. 
Donald M. Britton Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch 
Irwin M. Brodo John M. Gillett Theodore Mosquin Dan Strickland 
William J. Cody C. Stuart Houston Eugene G. Munroe John B. Theberge 
Francis R. Cook George F. Ledingham Robert W. Nero Sheila Thomson 
Ellaine Dickson 

2005 Council 
President: Mike Murphy Ronald E. Bedford —_ Diane Kitching Louise Schwartz 
Vice-President: Gillian Marston Fenja Brodo Karen McLachalan Hamilton David Smythe 
Recording Secretary: Susan Laurie-Bourque William J. Cody David Hobden Henry Steger 
Treasurer: Frank Pope Kathy Conlan Cendrine Huemer Chris Traynor 
Past President: Gary McNulty Francis R. Cook Diane Lepage Eleanor Zurbrigg 


Stanley Rosenbaum 


To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, : 
Westgate P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc @achilles.net. 
For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca 


The Canadian Field-Naturalist 


The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in — 
this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. 


PAP Registration Number 9477. Canada We acknowledge the financial support of the Government of Canada through the : 
Publication Assistance Program (PAP) toward our mailing costs. 


Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: cfn@ofne.ca 
Copy Editor: Elizabeth Morton 

Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2; (613) 759-1374 
Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5, e-mail: roy.john@pwegsc.gc.ca 


Associate Editors: Robert R. Anderson Paul M. Catling David Nagorsen 
Charles D. Bird Brian W. Coad Donald F. McAlpine 
Robert R. Campbell Anthony J. Erskine William O. Pruitt, Jr. 


Chairman, Publications Committee: Ronald E. Bedford 


All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal 
mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor. 


Subscriptions and Membership 

Subscription rates for individuals are $28 per calendar year. Libraries and other institutions may subscribe at the rate of $45 per 
year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $28 (individual) $30 (family) $50 (sustaining) and 
$500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) | 
must add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and 
Local Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to 
Libraries at $28 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies 
should be mailed to: The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post 
Publications Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: April-June 2005 (June 2006). 


Cover: A male Pardosa hyperborea, a small wolf spider charateristic of woods, barrens, and bogs of Newfoundland. This indi- 
vidual caught at Red Cliffe, Bonavista Bay, Newfoundland 3 August 2004 and photographed in the laboratory by Roy 
Ficken. See “An annotated list of the Spiders of Newfoundland” by J. R. Pickavance and C. D. Dondale pages 254-275. 


MCZ 
LIBRARY 


JUL 11 2006 


ARVARD 
April—June 2005 


The Canadian Field-Naturalist 


Volume 119, Number 2 


The Influence of Thermal Protection on Winter Den Selection by 
Porcupines, Erethizon dorsatum, 1n Second-Growth Conifer Forests 


ToppD N. ZIMMERLING 


Gartner Lee Limited, Suite N195, 3015-5" Avenue NE, Calgary, Alberta T2A 6T2 Canada 


Zimmerling, Todd N. 2005. The influence of thermal protection on winter den site selection by Porcupines, Erethizon dor- 
satum, in second-growth conifer forests. Canadian Field-Naturalist 119(2): 159-163. 


I investigated den type selection by Porcupines (Erethizon dorsatum), in relation to the thermal cover provided by the den type, 
over a period of four winters. Porcupines used log dens, stump dens and rock dens in proportion to the thermal cover provided 
by each den type. Based on behavioural observations of Porcupines, I assumed that the lower critical temperature for porcupines 
in my study area was -4°C. Both stump and rock dens provided adequate thermal protection, under most ambient conditions, 
to allow Porcupines to maintain their body temperature, without increasing basal metabolic rate. In most cases rock and stump 
dens maintained den temperatures above -4°C until ambient temperatures reached -12°C or lower. In contrast log dens pro- 
vided poor thermal protection, even in years of thick snowcover. When ambient temperatures dropped below -4°C, den tem- 
peratures within log dens were also recorded below -4°C. Log dens were used least often by Porcupines, whereas stump and 
rock dens were used most often. Despite the large number of potential dens available to Porcupines within the study area, den 
use was generally limited to three dens per porcupine per winter. The limited use of dens by an individual porcupine during 
winter may be related to the energetic cost of finding a new den or it may be related to specific selection criteria used by 
Porcupines. 


Key Words: Porcupines, Erethizon dorsatum, winter, lower critical temperature, temperature differential, winter den, den 


selection, den sharing, thermal cover, thermal protection, British Columbia. 


Cold temperatures can have dramatic impacts on 
mammals, including increasing individual mass loss 
and mortality rates (Moore and Kennedy 1985; Saether 
and Graven 1988; Sweitzer and Berger 1993). Despite 
the extreme cold weather conditions that may be en- 
countered throughout the northern portions of the Por- 
cupine’s (Erethizon dorsatum) range, the Porcupine 
appears to be poorly adapted for survival in colder cli- 
mates. Its dorsal pelage is sparse in comparison to 
other northern furbearers, and the soles of its feet are 
hairless and large (Folk 1966). Under winter conditions 
the lower critical temperature (ambient temperature 
at which the Porcupine must increase its basal meta- 
bolic rate above resting rate to maintain body tempera- 
ture) for Porcupines is estimated to be between -12°C 
(Irving et al. 1955) and -4°C (Clarke 1969), which is 
high compared to other mammals living in northern 
climates (Scholander et al. 1950a, 1950b; Irving et 
al.1955), 

Given that Porcupines have a high, lower critical 
temperature, cold ambient temperatures must consti- 
tute a major energy cost for them. To survive in cold 
temperature conditions the Porcupine has evolved a 
number of physiological and behavioural adaptations 
(Clarke and Brander 1973; Roze 1987; DeMatteo and 


Harlow 1997). One of the behavioural adaptations is 
the use of a winter den. Sweitzer and Berger (1993) and 
Roze (1989) found that Porcupines alter their foraging 
patterns in response to cold weather and spend more 
time in dens. Radiometric measurements (Clarke and 
Brander 1973) indicate that cover above the back of a 
Porcupine, such as a conifer branch, reduces radiative 
heat loss, and presumably a winter den (not measured 
in the Clarke and Brander study) provides even greater 
thermal protection than a conifer branch. 

The objective of this study was to determine the rel- 
ative thermal protection provided to a Porcupine by 
different den structures and to examine whether Por- 
cupines choose den types with respect to the thermal 
protection provided. Den switching and den sharing by 
Porcupines was also examined to compare winter den- 
ning behaviour of this study population with those from 
other areas. 


Study Area 

This study took place near Terrace, British Colum- 
bia, Canada (54°35', 12°42') in the Shames Valley 
(approximately 30 km west of Terrace; Figure 1). The 
study site was composed of three second-growth stands, 
(20-25 years old) located in the Coastal Western Hem- 


32 


160 


Terrace 


ra) 
\\ 


Vancouver 


FIGURE 1: Study site location within British Columbia, Canada. 


lock, wet submaritime (CWH,,.) biogeoclimatic sub- 
zone (Banner et al. 1993). The second-growth forest 
consisted primarily of Western Hemlock (Tsuga het- 
erophylla), Amabilis Fir (Abies amabilis), and Sitka 
Spruce (Picea sitchensis) with a minor component of 
Western Red Cedar (Thuja plicata). The study stands 
were located on a west-facing slope with gradient up to 
40% present. The lower boundaries of the study stands 
were at approximately 200 m elevation and the upper 
boundaries were at approximately 450 m elevation. 
Average minimum monthly temperatures varied con- 
siderably from month-to-month and from year-to-year 
over the four winters of the project. Fourteen of the 
20 months of this study had average minimum temper- 
atures of -4°C or lower and 6 of the 20 months had 
average minimum temperatures of —12°C or lower. 


Methods 

During the winters (November — March) of 1996 
to 2000, using systematic searches of the study area, 
33 Porcupines were located and captured. Once located, 
the Porcupine was netted, using a large dip-net, if the 
animal was out of its den. If the animal was in its den, 
a Tomahawk, single door live-trap (40 x 40 x 100 cm) 
was placed at the den entrance and left overnight. 
Once captured, the animal was drugged using a 10:1, 
ketamine (10 mg/kg): xylazine (1 mg/kg) combination. 
To reduce the time to recovery of smaller animals, 1 kg 
was subtracted from body weight when determining 
dosage. This procedure ensured that smaller animals, 
that could be processed more quickly and were more 
susceptible to exposure to cold temperatures, recovered 
more quickly than larger animals that required more 
processing time and were less susceptible to exposure 
to cold temperatures. 

Throughout the winter months from 1996 to 2000 
each collared animal was located three times per week, 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


using handheld telemetry. When an animal was found 
in a den the location was recorded using a Trimble 
ProXL GPS unit with differential correction (accurate 
to + | m), and a temperature measurement from inside 
and outside the den was made. The temperature was 
measured using a Barigo digital indoor/outdoor ther- 
mometer (accurate to + 1°C) with a 3 m probe cable. 
The probe was taped to heavy electrical wiring to give 
stiffness and to allow manipulation into the Porcupine 
den. When the Porcupine could be seen within the den, 
the probe was placed beside (but not touching) the 
animal. When the animal could not be seen the probe 
was inserted to a maximum of 2 m. In all cases the 
probe was positioned so as not to be resting on the 
den floor. The probe was left for 10 minutes to gain a 
stable temperature reading. Once the den temperature 
had been recorded the probe was then removed and 
placed at the top of snowcover, in a forest opening 
closest to the den site, to record ambient temperature. 

Although a single Porcupine may have used the same 
den over several weeks during the winter or during two 
or more winters, only a single temperature measurement 
was taken from each den site over the course of this 
study. In addition to the den use by collared Porcupines, 
den use by uncollared Porcupines was also noted (the 
type of den used) when discovered fortuitously. Tem- 
peratures were not recorded at these den sites unless it 
was confirmed (through visual observation) that the 
Porcupine was occupying the den at the time. 

To estimate the average maximum and minimum 
temperature within the study area, three monitoring sites 
were established within the study stand. Each moni- 
toring station had one max/min thermometer (Taylor 
indoor/outdoor maximum/minimum mercury thermo- 
meter) at snow level attached to the north side of a tree. 
These locations were visited once per week throughout 
the winter months and the maximum and minimum 
temperature for the week was recorded and averaged 
with the three readings. A mean monthly maximum and 
minimum was then calculated for each winter month 
using the weekly readings. 


Statistical Analysis 

I compared the thermal protection provided by each 
den type using a single factor analysis of variance of 
the temperature differentials for each den type. Where 
required a Tukey test was used to determine which 
means were different (Zar 1974). Temperature differ- 
entials were calculated by subtracting the ambient tem- 
perature from the den temperature to give a positive 
number. When den temperatures were lower than ambi- 
ent temperatures the differential was negative. 


Results 

Porcupines within the Shames Valley study area used 
four distinct den types: (1) log dens, which were defined 
as dens within fallen, hollow logs or dens located under 
a fallen tree (usually having been created by trees that 
had been cut down during commercial harvesting, but 


2005 


left on-site). (2) Stump dens, which were defined as dens 
located beneath the stump of a previously harvested 
tree. In all cases these dens were located in the rootwad 
of the stump, and not in any kind of hole within the 
stump itself. (3) Rock dens, which were dens located 
within the crevices of rock outcroppings. These dens 
were usually very deep and often had multiple, potential 
entrances. (4) Pre-excavated dens, which were rare in 
the study area, consisted exclusively of dens excavat- 
ed by Coyotes (Canis latrans) into soft soils on gully 
slopes. 

A total of 46 dens were measured to determine the 
temperature differential between den and ambient tem- 
peratures. Unfortunately, owing to the low number of 
pre-excavated dens used, no measurements for this den 
type were obtained. Rock and stump dens provided 
almost identical thermal protection, with mean temper- 
ature differentials of +5.4°C and +5.2 °C, respectively 
(Figure 2). Log dens provided the poorest insulation 
for Porcupines, as the best temperature differential re- 
corded for a log den was +4°C and the mean was +1.3°C 
(Figure 2). This compares with the best temperature 
differential for a rock den of +9°C and a stump den of 
+10°C. Analysis of variance, with the associated Tukey 
test, showed log dens provide significantly less thermal 
protection than both rock and stump dens (ANOVA p = 
A xnOfod f= 2,43: R= 9.42). 

In the single case where a log den was measured 
while ambient temperature was below -12°C, the den 
temperature was also below -12°C (-16°C ambient, 
-15.3°C den). In nine cases where den temperatures were 
recorded in stump and rock dens while ambient tem- 
peratures ranged from -12°C to -20°C, the den temper- 
atures remained above -12°C. 

Of 24 observations of Porcupines out of their dens 
during the day, only one occurred when the ambient 
temperature was below -4°C (temperature was -5°C). 
This observation involved a Porcupine moving down 
a tree and travelling to a stump den, which it entered. 
All other observations of active animals involved Por- 
cupines feeding or resting in trees when temperatures 
were between -2°C and +6°C. 

If -4°C is the lower critical temperature for Porcu- 
pines, then log den temperatures were at or below the 
lower critical temperature in all eight cases where 
ambient temperature was below -4°C. Rock den tem- 
peratures were at or below -4°C in four out of twelve 
cases where ambient temperature was below -4°C, and 
stump den temperatures were at or below -4°C when 
ambient temperatures were below -4°C in four out of 
fifteen instances. In the cases where stump and rock den 
temperatures were below -4°C, the ambient temperature 
was -12°C or colder and the level of thermal protection 
provided by the den was high (+6.7°C to +10.0°C 
temperature differential). 

The use of pre-excavated and log dens was similar 
from year-to-year, with pre-excavated dens being used 
in only one year (1996-1997) and log dens being used 
infrequently in all years (Table 1). In three of the four 


ZIMMERLING: WINTER DEN SELECTION BY PORCUPINES 


161 


a 


Mean Temperature Differential 
> 


FIGURE 2: Mean temperature differential (den temperature — 
ambient temperature in °C) (+ | s.d.) of winter den 
types used by Porcupines in the Shames Valley (All 
data from 1996 to 2000 combined). 


years the majority of dens used were stump and rock 
dens, the two den types with the best temperature dif- 
ferentials. The biggest single change in den use, from 
year-to-year was in the use of stump dens in 1997-1998. 
During this winter, the use of stump dens dropped to 
the lowest usage of the study (13.8%), while the use 
of rock dens increased to the highest usage (69.0%) 
(Table 1). The winter of 1997-1998, based on aver- 
age weekly temperatures, was the warmest winter of 
the four-year study (Table 2). January was the only 
month in which 1997-1998 did not have the warmest 
maximum and minimum average temperatures, com- 
pared to the three other study years. 

The number of dens used by a single Porcupine in 
one winter was constant over three winters of the study. 
In 1996-1997, 1998-1999 and 1999-2000 individual 
Porcupines used an average of 3.4 dens/Porcupine, 
3.3 dens/Porcupine and 3.3 dens/Porcupine, respective- 
ly. The average number of dens used by a Porcupine 
dropped in 1997-1998 to 2.1. 

Throughout the four winters of this study individual 
Porcupines did not always use the same den sites from 
year-to-year. In some cases new dens were used every 
winter and in a small number of instances the same den 
was used by different Porcupines in two different years. 
Only one instance of two Porcupines using the same 
den at the same time was noted. This den sharing oc- 
curred in the winter of 1998-1999 and involved an adult 
female and a juvenile female. It was unknown if the 
two animals were related, but they shared the same den 
for 22 days without any indication of disputes occurring 
within the den (no sounds when den was observed by 
researchers). 


Discussion 

Porcupines in this study used four distinct den types 
(log, stump, rock or previously excavated dens); how- 
ever, previously excavated dens were only used to a 
minor degree during a single winter. Unlike the results 
reported by Roze (1987), Griesemer et al. (1994), and 
Griesemer et al. (1996) all animals in this study used 


162 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |: Winter den type selection by Porcupines in the Shames Valley (1996-2000). 


Winter n Log 

1996-1997 47 17.0% 
1997-1998 20 17.2% 
1998-1999 24 12.5% 
1999-2000 23 17.4% 
Total 123 16.3% 


dens extensively throughout the winter and although 
roosting in trees during the day was noted, none spent 
more than a day in the shelter of conifer trees rather 
than in a den. 

The relative use of the three common den types mur- 
rored closely the temperature differentials found be- 
tween them. Rock and stump dens had the largest 
temperature differentials and were used equally often 
through most of the study. An exception to this was the 
winter of 1997-1998, when rock dens were used more 
than in any other year and stump dens were used less 
often. This winter was also the warmest winter of the 
study. The warmer average temperature resulted in more 
mid-winter snowmelt than in other study years, and 
this may have influenced den choice by Porcupines. 
Stump dens tend to be low-lying and are susceptible to 
water seepage. As a result, increased snowmelt may 
have resulted in water in the den and thus lower use 
by Porcupines. 

Log dens had the poorest temperature differential and 
were used least often among the three common den 
types. Even in years of increased snowfall, when the 
thick snowpack would be expected to provide increased 
thermal cover to a log, Porcupines did not increase the 
use of this den type. In all years, when ambient tem- 
peratures dropped below -4°C, log dens did not pro- 
vide adequate thermal protection to Porcupines. 

Roze (1989) found similar results in den use by 
Porcupines in the Catskill Mountains of New York 
State. Rock crevices were the most used denning struc- 
ture (70% of dens) and hollow logs were least often 
used (< 10% of dens). A key difference between the 
Porcupines in the Catskill Mountains and the Porcu- 
pines in the Shames Valley is the use of stump dens 
and standing, hollow trees. In the study area chosen 
by Roze (1989), Porcupines used standing trees with 
hollow openings for dens 20% of the time, and never 
used stump dens, as seen in my study. The difference 
in den use is related to forest structure. My study site is 
a second-growth forest, 20—25 years of age, and, as a 
result, there are no trees within the study area that are 
old enough to have developed hollow openings in the 
stem. The high use of stump dens is related to previ- 
ous logging activities. The removal of the large old- 
growth trees resulted in large stumps being left behind. 
As the root structure of these stumps begins to decay 
a natural opening is created beneath the stump in the 


Den Type 
Stump Rock Pre-excavated 
38.3% 38.3% 6.4% 
13.8% 69.0% 0.0% 
50.0% 37.5% 0.0% 
52.2% 30.4% 0.0% 
37.4% 43.9% 2.4% 


root wad. Although Roze’s study area encompassed an 
abandoned farm, he mentions no evidence of recent 
(in the last 30 years) logging on the site. As a result, 
stumps would only be created through natural mortality 
of trees. Speer and Dilworth (1978) also reported that 
the majority of den sites in their New Brunswick study 
site were located in the roots of trees. These dens, how- 
ever, were not under stumps, but located in the roots 
of wind-thrown trees. Only one of 69 dens was located 
in a standing, hollow tree. 

Based on behavioural observations of Porcupines 
during my study, I assumed that the lower critical tem- 
perature for Porcupines in the study area is closer to 
—4°C as reported by Clarke (1969), rather than the 
—12°C reported by Irving et al. (1955). In my study 
area, under most winter temperature conditions, Por- 
cupines are able to rely solely on the thermal cover 
provided by a rock or stump den to reduce the energy 
requirements for maintaining body temperature, as 
den temperatures are maintained above -4°C. When 
ambient temperatures drop below —12°C, it appears 
Porcupines are forced to increase their metabolic rate 
to maintain body temperature within rock and stump 
dens, however, on average, temperatures within the 
study area were above —12°C. 

Porcupines in my study were usually solitary in their 
den occupancy, but there was one instance where two 
Porcupines used the same den at the same time. The 
sharing of dens by Porcupines has been reported by 
Roze (1987), who found 12% of dens to be occupied 
by two Porcupines (usually a male-female pair). Dodge 
(1967) also described den sharing by Porcupines in 
western Massachusetts. In contrast, Dodge and Barnes 
(1975), Brander (1973) and Shapiro (1949) found that 
Porcupines rarely shared dens in their studies. The oc- 
currence of winter den sharing appears to be directly 
correlated with the abundance of den sites in a partic- 
ular area. All three studies where Porcupines were 
found to rarely share dens were described as having 
an abundance of den sites, whereas Roze (1987) de- 
scribes both his study site and that of Dodge (1967) 
as having limited den sites. Griesemer et al. (1996) 
when comparing den sharing in two areas with differ- 
ent den availability in Massachusetts found that avail- 
ability of den sites did influence den sharing by Por- 
cupines. The results from my study also support this 
argument as the Shames Valley has a large number of 


2005 


potential den sites (as indicated by numerous unoc- 
cupied, but previously used dens). 

Despite the large number of available dens, Porcu- 
pines in the Shames Valley used only a few dens per 
individual during a single winter (approximately three 
dens/Porcupine during 1996-1997, 1998-1999, and 
1999-2000). In the warm winter of 1997-1998 the aver- 
age number of dens used per Porcupine dropped to two. 
Increased snowmelt in this year may have reduced 
the number of dens available to Porcupines, owing to 
water seeping into den sites. Alternatively, the warmer 
temperatures may have also increased the energetic 
cost of travel for Porcupines by creating a soft snow- 
pack, thereby restricting Porcupine movements. Roze 
(1989) reported a similar impact, with high snowfall 
events reducing Porcupine winter movements and 
numbers of dens used. 

Porcupines choose den structures in relation to the 
relative thermal cover provided by the structure. In 
the case of the Shames Valley, Porcupines are taking 
advantage of an abundance of stump dens, which have 
been created as a result of past commercial forest 
harvesting in the area. Given the role of the winter den 
in Porcupine winter ecology (Roze 1987, 1989; Griese- 
mer et al. 1994; Zimmerling and Croft 2001) it is like- 
ly that the creation of dens with high thermal cover is 
increasing the areas over which Porcupines can main- 
tain winter ranges. Without the past forest harvesting 
Porcupines in the Shames Valley would have been lim- 
ited to rock dens. Consequently Porcupine den locations 
would be more confined to steep areas where rock has 
been exposed. With the past forest harvesting activi- 
ties Porcupines can find dens with good thermal cover 
throughout the second-growth stand and can establish 
winter ranges accordingly. 


Acknowledgments 

This research was supported financially by Skeena 
Cellulose Inc., Terrace Operations, and the British 
Columbia Ministry of Forests, Kalum Forest District. 
Major funding was supplied by Forest Renewal British 
Columbia. | 

I thank M. D. Bahr, M. A. Bahr, A. Coosemans, W. 
Croft, C. Croft, M. Dawson, B. Delahanty, M. Hamil- 
ton, C. Jardine, K. Kranrod, P. Kranrod, M. Krisinger, 
J. Pylon, W. Sheridan, P. Skinner, L. Zimmerling, and 
R. Zimmerling for their hard work under extreme 
field conditions. Thanks to L. Zimmerling and C. J. 
Krebs for reviewing an earlier draft. 


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Denning patterns of porcupines, Erethizon dorsatum. Cana- 
dian Field-Naturalist 110: 634-637. 

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Received 4 March 2002 
Accepted 14 May 2005 


Black Bear, Ursus americanus, Ecology on the Northeast Coast 
of Labrador 


KEITH CHAULK!, SOREN BONDRUP-NIELSEN2, and FRED HARRINGTON? 


' Box 382, Station “C”, Goose Bay, Labrador, Newfoundland and Labrador AOP 1E0 Canada 
* Acadia University, Wolfville, Nova Scotia BOJ 1X0 Canada 
> Mount Saint Vincent University, Halifax, Nova Scotia B3M 2J6 Canada 


Chaulk, Keith, Soren Bondrup-Nielsen, and Fred Harrington. 2005. Black Bear, Ursus americanus, ecology on the northeast 
coast of Labrador. Canadian Field-Naturalist 119(2): 164-174. 


Twenty-three Black Bears (Ursus americanus) were captured, 20 were measured, marked and/or radio collared, in north- 
eastern Labrador, between 1996 and 1997. Bears used sea ice for travel, coastal islands for denning, hunted adult Caribou 
(Rangifer tarandus), and were the possible cause of Moose (Alces alces) calf mortality. Body sizes were small, median 
weight of adult females was 48 kg, and the sex ratio for captured subjects was 1:1. Four of six radio-collared females gave 
birth during the winter of 1997, female reproductive histories suggest delayed sexual maturity. Den entry occurred between 
October and December 1996; spring emergence occurred between April and May 1997, with estimated denning period 
ranging from 148-222 days. Visual observations of habitat use by radio collared subjects (n = 10) were not tested statistical- 
ly but suggest that barren areas are used nearly as much as forest. Location data from three GPS collars deployed on three 
adult females were analysed using Chi-square goodness-of-fit test with Bonferroni correction; two females appeared to pre- 
fer forest habitats (p < 0.05). 


Key Words: Black Bear, Ursus americanus, Labrador, telemetry, home range, morphology, denning, habitat, translocation, 


activity. 


Although the Black Bear (Ursus americanus) has 
been studied throughout most of its range, it had been 
virtually ignored in Labrador. This changed in 1989, 
when a five-year research program was initiated to 
document Black Bear ecology near Hebron Fiord in 
northern Labrador (Veitch 1992; Veitch 1994; Veitch 
and Harrington 1996). The Hebron bears demonstrat- 
ed atypical characteristics such as heavy reliance on 
animal protein, home ranges orders of magnitude larg- 
er than bears to the south, and year-round activity on 
barren habitats, earning them the name “‘barren-ground 
Black Bear” (Veitch 1992). However, it remained un- 
clear if bears in more southerly regions of Labrador 
shared these barren-ground characteristics, if they dis- 
played characteristics more closely associated with 
forest dwelling bears or were intermediary between 
the two. 

Shortly after the completion of the Hebron study in 
1993, a large nickel deposit was discovered approxi- 
mately 200 km to the south, near Voisey’s Bay, Lab- 
rador. The anticipated development of that deposit, 
and the associated environmental impact assessment 
prompted the present study on the ecology of Black 
Bears in the Voisey’s Bay area. This article describes 
observations made during 1996 and 1997 regarding 
Black Bear habitat use, den site characteristics, den- 
ning period, food habits, daily activity, morphology, 
productivity, and demographics. 


Study Area 
The study area (Figure 1), approximately 1600 km/?, 
located south of Nain, Labrador, is rugged with ele- 


vations ranging from 0 to 650 m above sea level (asl). 
The western section is relatively flat, with the main 
habitat types being heath and rock barrens. The central 
region is comprised of rounded topography, valleys, 
and depressions and the main habitat types include 
heath and rock barrens. The eastern portion is charac- 
terized by low-lying coastline, sheltered river valleys, 
and rolling hills (JWEL 1997*). The habitat mapping 
component of this study was defined by a 1:20 000 
ecological land classification (SWEL 1997*) and com- 
prised approximately 364 km? of the larger study area 
(Figure 1). 

Weather and climate information collected by Envi- 
ronment Canada at Nain from 1951 to 1989 show that 
the mean monthly temperature varies from -19°C in 
January to 10°C in July, with a mean annual tempera- 
ture of -3°C. Mean annual precipitation is 740 mm, 
with the highest monthly rainfall recorded during July 
(79 mm) and maximum monthly snowfall occurring 
in January (87 mm). Snow and ice can persist until 
July (Environment Canada 1989). 


Methods 

Human activity related to mineral exploration in the 
study area at the time of this study, although recent, 
was at an all time high, and bear-human conflicts were 
common. To reduce the number of bears killed, camp 
personnel initiated a Black Bear translocation pro- 
gram. In addition the study team used leg snares, and 
darting from a helicopter to capture Black Bears. All 
bears captured in culvert traps were moved away from 
project activities by helicopter: translocation distances 


164 


2005 


varied. For most translocations, a second helicopter 
was used to transport the study team to the release 
location. All marked Black Bears were initially tran- 
quilized with 4-7 mg/kg of Telazol (White et al. 1996). 
All subjects were monitored during recovery from 
anesthetic and were revisited 24-30 hours after being 
tranquilized. 

Most bears were aged, weighed, and sexed and signs 
of estrus (Coy and Garshelis 1992) were recorded. The 
senior author supervised all measurements in order to 
reduce researcher variation (Eason et al. 1996). Bears 
were marked with Flex-Lok plastic ear tags (Ketchum 
Manufacturing, Ottawa, Ontario). A premolar was ex- 
tracted from each bear and sent to Matson’s Labora- 
tory (Montana) for cementum annuli analysis (Dim- 
mick and Pelton 1996), which was used to determine 
age. For purposes of analysis and discussion, bears 
were classed as adults if they were 5 years and older, 
and sub-adults if 3-4 years of age; cubs (ages 1-2 
years) were not captured or marked. The alpha-numeric 
code used to identify each bear was formatted as fol- 
lows: age class (1.e., A,S,U), sex (i.e., M,F,U), capture 
sequence number (i.e., AMBO1 = adult male bear 
number 01; SFB16 = sub-adult female bear number 16; 
UUB22 = Unknown sex and age bear number 22). 

Seven VHF collars (Holohil Inc., Carp, Ontario) 
and three Global Positioning System (GPS) collars 
(Lotek Engineering Inc., Newmarket, Ontario), were 
fitted on Black Bears during June and July 1996. 
GPS collars weighed 1.36 kg and recorded geodetic 
coordinates (latitude and longitude, WGS 84), tem- 
perature, time, date, fix status, horizontal dilution of 
precision (HDOP), convergence (distribution of satel- 
lites above the horizon), and activity every three hours 
during a total of eight fix attempts/day. 

Activity was measured by means of mercury switch 
activations; these were summed every ten minutes and 
averaged every three hours to provide a single activity 
count at the time of data logging. Activity data were 
summarized and expressed.as a function of an average 
day relative to sunrise and sunset. After failure, GPS 
collars were retrieved and replaced with VHF collars. 
Two-dimensional locations were assumed to have an 
error radius of 50 m (Lotek Engineering Inc., 1996*); 
differentially corrected GPS data were assumed to 
have an error radius of 10 m (Moen et al. 1996); only 
three-dimensional (3-D) fixes were differentially cor- 
rected. 

Telemetry flights were conducted between 24 June 
and 15 October 1996 using a Bell 206B helicopter. 
Once a radio signal was localized to a small area 
(100 m radius), an effort was made to acquire a visual 
fix. When a visual fix was not possible, the subject’s 
location was estimated. The helicopter’s GPS was used 
to record the location of all observations. Ground 
telemetry was conducted during the same period when 
aerial telemetry or animal handling were not in pro- 
gress. GPS locations obtained from the helicopter and 


CHAULK, BONDRUP-NIELSEN, AND HARRINGTON: BLACK BEAR ECOLOGY 


165 


oy Hebron 
S 


Labrador 


Labrador Sea 


_ Nain 


~ {/ 1/111] Land Classification 


: *. —————] Study Area 
FiGurE |. Black Bear Study Area Voisey’s Bay Labrador 1996 


and 1997. 


handheld GPS units were not differentially corrected 
and were assumed to have an accuracy of +/-100 m 
(Moen et al. 1996); geodetic data were recorded in 
World Geographic System (WGS) 84 latitude and 
longitude, and were converted to latitude and longi- 
tude, North American Datum 83 using MAPINFO. 

We searched for den sites in October and November 
1996, while conducting ground and aerial telemetry. 
Frequency of monitoring flights was reduced after 
October. Monitoring flights were conducted on 3, 4 
and 29 November 1996. Known Black Bear dens were 
monitored for activity on a monthly basis from Janu- 
ary to March 1997. In the spring of 1997 four aerial 
monitoring sessions were conducted to determine the 
timing of den emergence (10 April, 26 and 27 April, 
25 May, and 12 June). Due to gaps in monitoring we 
were not able to determine exact dates for either den 
entry or emergence. For example, the maximum date 
of den entry was the date the study team was able to 
confirm if a bear had entered its den. The minimum 
date of den entry is the date of the prior survey when 
the bear was still active. The converse was done for 
den emergence. We assumed each bear entered/exited 
its den halfway between the minimum and maximum 
dates of den entry/emergence. However, for ease of 
calculation the estimated den occupancy as the differ- 
ence (in days) between maximum date of den entry in 
1996 and the maximum date of den emergence in 1997. 

Den sites were assigned to three habitat types: for- 
est, barren, and other based on visual inspection of the 
surrounding landscape. Forest habitat was any area with 
canopy height exceeding | m, barren habitat was any 
non-wetland area with less than 5% canopy cover. 
Visual observations of radio collared Black Bear habi- 
tat use was recorded in the field, and habitats were 
classed using the same criteria as den site habitat class- 
ification. Visual observations reported in this study were 
not tested for selection because many of the observations 
occurred outside the area described by the 1:20 000 
digital vegetation maps, and therefore habitat availabil- 
ity could not be calculated. 


166 


As part of the environmental baseline research for 
the Voisey’s Bay Environmental Impact Statement 
(EIS), digital habitat maps (1:20 000) were developed 
based on interpretation of aerial photography and ground 
surveys (SWEL 1997*); 21 habitat classes were differ- 
entiated. However, given the limited number of loca- 
tions per subject in each habitat type, we grouped these 
21 habitat classes into three broad categories: barren, 
forest or other. 

For the three GPS collared bears, Minimum Convex 
Polygons (MCP) and habitat maps were used to esti- 
mate habitat availability. Observed versus expected 
habitat use was tested for evidence of habitat selection 
using Chi-square goodness of fit (Neu et al. 1974). If 
the test was significant at p < 0.05, availability was 
compared to the 95% confidence interval (CI) for that 
habitat. The 95% CI was based on the normal approx1- 
mation to the binomial distribution, with a Bonferroni 
correction for multiple testing (Neu et al. 1974). 
Throughout the text + is used to represent standard 
deviation, whereas in the tables it is denoted by SD. 


Results 

There were 44 capture events (23 bears, 20 of these 
were marked, and 11 were captured on more than one 
occasion) between June and November 1996 (Table 1). 
Sex and age were not determined for three subjects, as 
camp officials captured and released these bear without 
participation by the study team. Twenty Black Bears 
were marked with ear tags and/or radio-collars, but no 
more than 10 Black Bears were fitted with radio-collars 
at any given time. Culvert traps and leg snares account- 
ed for 27 and 9 captures respectively, one bear was 
darted from helicopter (Table 1). 

Of the 20 marked bears, 10 were male and 10 were 
female. Of the 22 bears that were assigned to an age 
class, 10 were sub-adult and 12 were adult; cementum 
analysis revealed ages from 2 to 23 years (Table 1). 
The mean cementum age of adult males was 12.5 + 
7.0 years (n = 4), the mean cementum age of adult 
females was 9.1 + 3.6 years (n = 8). The mean age of 
sub-adults was 2.8 + 0.8 year (n = 5). The mean age of 
all males was 8.1 + 7.4 years (n = 10), and the mean 
age for all females was 8.0 + 3.0 years (n = 10). Thir- 
ty percent of bears were 10 years or older, and 67% 
were older than 6 years (n = 17). AMB18 was radio 
tracked and observed in spring of 1997, making it 
24-years-old at time of last contact (Table 1). 

Two of 10 females were in estrus at the time of 
capture (AFBO2 and AFB04). Two opposite-sex pairs 
of Black Bears were observed together for extended 
periods during July 1996. Only one collared bear 
(AFBO07) was actively caring for cubs during 1996; this 
family group was still intact at the time of den emer- 
gence in 1997. According to reproductive histories 
based on cementum analysis, only AFB06 had given 
birth prior to 1996, at ages 6, 9, and 11 years; AFBO6 
also produced cubs for the fourth time at the age of 14 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


during the winter of 1997. Radio tracking during the 
spring of 1997 showed that four of the five collared 
females gave birth during the winter of 1997; the mean 
age at first reproduction was 8 + 1.6 years (range = 6 
— 10 years). (Table 2). 

Based on analysis of cementum annuli two females, 
AFBO03 and AFBO04, had not reproduced at the time 
of capture (age 7 and 6, respectively). Five of seven 
females (71%) produced cubs over a two-year period. 
From 1996 to 1997, 11 cubs of the year (COY) were 
produced, mean = 1.8 + 1.0 cubs/adult female, n = 6 
(Table 2), the mean annual rate of reproduction = 0.8 
+ 0.6 cubs/female/year (range 0-1.5, n = 7). 

Fifteen bears were translocated a total of 25 times 
(Table 3). Estimated return periods from translocation 
point to capture site for 11 bears ranged from 1-55 days 
(mean = 18.3 + 19.0 days); estimated rate of return 
ranged from 0.5 km/day to 6.5 km/day with mean rate 
of return of 2.6 + 2.2 km/day (Table 3). Four bears in 
the study area were shot by mining camp and provin- 
cial wildlife officials in 1996: two were collared 
(AMBO1 and SMB11), one was tagged (AMB19), and 
one was unmarked (UMB17). Three bears died during 
1997: SFB16 (natural causes), SMB 13 (shot by hunter), 
and SMB10 (shot by camp personnel) (Table 1). Six 
of seven known mortalities were males (three adults, 
two sub-adults, one age unknown). A post-mortem 
examination of AMB19 at the Atlantic Veterinary Col- 
lege (University of Prince Edward Island, Charlotte- 
town) showed that this bear had previously been shot 
and had been suffering from lead poisoning for ap- 
proximately six weeks prior to its death. No Black Bears 
died as a result of capture or handling by the study 
team. 

Between June and October 1996, weights and phys- 
ical measurements were recorded for 20 adult and 
sub-adult bears. Weights for all subjects ranged from 
27 to 130 kg (n = 20); adult males (120 + 19 kg, n= 4) 
were about twice as heavy as females (48 + 13 kg, 
n = 8). The heaviest male and female were 8 and 7 
years old, respectively. Chest measurements for all sub- 
jects ranged from 64 to 112 cm (n = 18) with adult 
males averaging 103 + 13 cm (n = 4) and adult fe- 
males 87 + 8 cm (n= 7). 

Activity counts from the GPS collars were averaged 
and graphed to represent mean daily activity. Minimum 
daily activity occurred 2-3 hours after sunset (23:00). 
Peak activity occurred 3-4 hours before sunset (17:00). 
Activity was relatively constant across other time inter- 
vals (Figure 2). 

Eighteen den sites were located during 1996: 7 in 
forest, 6 in barren and 5 in shrub thicket (Table 4). The 
entrance to all dens faced south or southwest. Eight 
dens were unoccupied and were located prior to the 
start of fall denning; 10 dens were located after den- 
ning. One bear left the mainland in late summer 1996 
and took up residence on a large coastal island where 
it excavated a den in a sandy spot underneath a shrub 


167 


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BLACK BEAR ECOLOGY 


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168 


TABLE 2. Summary of Cubs of the Year (COY), Reproductive History and Age at first Birth for marked adult female Black Bears in Voisey’s Bay, Labrador study area during 1996 and 


1997. Age at first Birth, and Reproductive History were discerned through analysis of cementum annuli (Coy and Garshelis 1992). 


Age at 
first birth 


Annual 


Rate (COY) 


Total 
Cc 


(COY) 


Age 
(96) 


ID 


Reproductive History 


1997 OY 


1996 


In estrus June (1996) 


N 


AFBO2 
AFBO3 
AFB04 
AFB06 
AFBO7 
AFBO8 
AFBO09 


Lost contact with this bear in June 1996 


In estrus June (1996); non-productive (1996 & 1997) 


Cubs at age 6, 9, 11, & 14 (1997) 


6 
13 


10 


N 


Two cubs at time of capture in June (1996) 


10 


185 


(oe) 


6 


N 


8 


Count 
Sum 


THE CANADIAN FIELD-NATURALIST 


40 


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Vol. 119 


thicket (Table 4). Three radio collared females entered 
dens by mid-October 1996. A mother with two cubs 
(AFBO7), left her den in late October and moved to a 
second den by early November. Six other bears (5 fe- 
males and | male) entered their dens by late Novem- 
ber, while one bear, AMB18 was still active on 29 Nov- 
ember 1996 (Table 4). In 1996, Black Bears were first 
sighted in March. Monitoring flights conducted in 
1997 found that 10 of 11 radio-collared bears had 
emerged from their dens between 27 April and 27 May, 
1997. The mean length of the estimated den period was 
190 days + 28.2 days (n = 9). 

A total of 185 visual observations were made of 
Black Bear habitat use based on aerial tracking of 
VHF collared subjects: subjects were observed in 
forest on average 54% + 21.5, barrens = 40% + 25.7, 
and other = 6% + 7 (Table 5). Individual bears varied in 
their relative use of barren and forest habitats. For 
example, 86% of AFB02’s locations (n = 28) were on 
barrens, whereas only 10% of AFBO08’s locations (n 
= 20) occurred in similar habitat (Table 5). 

Three adult female Black Bears were tracked using 
GPS collars during July and August, 1996 (n = 24, 45 
and 56 locations, respectively). Bear locations most 
commonly occurred in forest habitats: Spruce/Fir/ 
Dwart-Shrub, Birch Thicket, Black Spruce Lichen, and 
Tuckamore. Chi-square analysis of gross habitat indi- 
cated that two bears (AFBO04 and AFBO8) occupied 
habitats differently from availability (p < 0.02); they 
appeared to prefer forest more than the other two pri- 
mary habitat types. The other subject (AFBO2) ap- 
peared to use all three habitats in accordance with 
availability (Table 6); forest habitat comprised 65% of 
her home range. During May and June the study team 
observed three incidences of Black Bears walking on 
sea ice up to 2 km from shore. 

Two incidences of ungulate predation/scavenging 
by Black Bears were observed during 1996. The first 
incident occurred in April and involved an unmarked 
adult male Black Bear attacking an adult female Cari- 
bou (Rangifer tarandus), which later died from its 
wounds and was partially eaten by the bear. The sec- 
ond incident occurred during an unsuccessful attempt 
to dart a Black Bear from helicopter in June 1996. The 
Black Bear was observed walking along an esker near 
a river. As we circled to dart the bear, it ran to the river 
and retrieved a dead Moose (Alces alces) calf and pro- 
ceeded to run with the carcass in its mouth for approx- 
imately 50 m. The bear then dropped the carcass and 
ran into the forest. An adult female Moose was seen 
within 200 m of the carcass. 


Discussion 

After the Muskox (Ovibos moschatus), Black Bears 
have the lowest reproductive rate of any land mammal 
in North America (Jonkel and Cowan 1971). Elowe and 
Dodge (1989) reported ages at first reproduction for a 
hunted population of Black Bears in Massachusetts; 
mean age at first reproduction was 3.7 + 0.7 years 


2005 

200 
> 
> 100 
17) 
<x 


2 5 8 11 
Time 


Sunrise 


CHAULK, BONDRUP-NIELSEN, AND HARRINGTON: BLACK BEAR ECOLOGY 


169 


14 i 1520 


Sunset 


23 


FiGuRE 2. Mercury Switch Activations (Activity) averaged over 8, three-hour time periods for three adult female Black 
Bears near Voisey’s Bay, Labrador. Observations were downloaded from data loggers housed in the GPS collars and 
occurred over 50-day period from 25 June to 13 August 1996 (n = 1080). The range of times for sunrise and sunset 
during this period are shown on the x-axis. On 25 June sunrise occurred at approximately 03:19 and set at 21:03 
Atlantic Daylight Savings Time (ADST); on 13 August sunrise occurred at 04:32 and set 19:53 (ADST), times for 
sunrise and sunset are based on an elevation of 400 m for the community of Nain, Labrador (Environment Canada 
1998). Dark squares indicate mean activity. Horizontal lines indicate the 95% Confidence Interval for the median. 


(range 3 — 5). Rogers (1993*) reported that the age at 
first reproduction for wild bears in eastern Minnesota 
was 6.3 years (n = 17). The mean age at first repro- 
duction for females in our study area was 8 years, 30% 
older than wild bears in Minnesota (Rogers 1993*) 
and 54% older than bears in Massachusetts (Elowe 
and Dodge 1989). 

Black Bear productivity in the Voisey’s Bay area was 
much higher than reported by Veitch and Harrington 
(1996) in northern Labrador. They tracked eight adult 
females for 22 bear years and during this time cubs 
accompanied females in 3 years (14%). In our study 
seven females were tracked for an equivalent of 13 bear 
years, and during this time cubs accompanied females 
in 5 years (38%). 

Due to logistical constraints we were not able to 
determine litter size at birth, so direct comparison with 
litter sizes in other areas of North America are difficult. 
We were able to document the number of cubs with 
each marked female during 1996 and spring of 1997. 
During this period 7 females produced 11 cubs (1.6 + 
1.1 cubs/female, n = 7); however, if we exclude fe- 
males that did not produce cubs the average number 
of cubs/female increases to 2.2 + 0.4 (n = 5). This 
compares to an average litter size = 2.4 cubs/female 


in Massachusetts (Elowe and Dodge 1989) and aver- 
age litter sizes ranging from 2.1 to 3.4 cubs/female in 
Minnesota, where litter size varied with foraging be- 
haviour and reproductive history (Rogers 1993*). 

We did not include observations of cubs associated 
with marked females in our capture statistics. In our 
study we found that sub-adults comprised 48% and 
adults 52%. Young and Ruff (1982) conducted a re- 
moval experiment on a population of Black Bears in 
central Alberta. For comparison purposes, we excluded 
cubs from their pre-removal capture data (1968-1971) 
and recalculated age structure statistics; sub-adults 
comprised 28% and adults 72% (n = 302). Kohn and 
Rolley (2000*) tabulated age structure data for bears 
harvested during 1998-1999 in Wisconsin. We re-cal- 
culated their age structure statistics with cubs exclud- 
ed; sub-adults comprised 54% and adults 46% (n = 
1664). Based on these comparisons the age structure 
of our study set was older than that reported in Wis- 
consin (Kohn and Rolley 2000*) and younger than that 
reported in Alberta (Young and Ruff 1982), albeit our 
sample sizes were very small in comparison. 

Six of 21 bears died during our study period trans- 
lating into a 28.6% mortality rate; all were male and 
five were sub-adult. No bears died as a result of handl- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE 3. Summary of Black Bear translocation and homing near Voisey’s Bay, Labrador during 1996. 


ID Captures Translocations Date 
SMBO0O0 3 3 06/19/96 
08/19/96 
10/08/96 
AMBOI? 3 l 08/02/96 
AFBO02? 3 l 07/29/96 
AFBO03 l 
AFB04? 2 
AMBO05 1 l 06/23/96 
AFB062 l iS 06/23/96 
AFBO07? I 
AFBO08? a 2 07/10/96 
08/22/96 
AFB09? Z 2 07/31/96 
08/11/96 
SMB 10? 2 2 07/31/96 
08/31/96 
SMBI1 I? Z. | 08/01/96 
AFB 12 4 4 08/01/96 
08/08/96 
08/18/96 
09/20/96 
SMB 13 I ] 08/02/96 
SFB 14 | 
SMB15 I 
SFB 16? Zz pa 08/26/96 
10/05/96 
SMB17 | 
AMB18 1 I 10/21/96 
AMB19 2 08/27/96 
10/18/96 
UUB20 I | 06/19/96 
Count 21 15 25 
Mean Ue) 7 08/13/96 
Median 2 l 08/08/96 
SD 1.0 0.9 a9 
Range 3 3 124 


Distance Return Rate of Return 
(Km) Period (days)! km/day 

| 1 1.0 

30 

26 

13 2 6:5 

31 6 3:2 

25 

12 22 0.5 

12 5 2.4 

23 8 2.9 

16 10 1.6 

56 

13 4 3.2 

52 

20 Bal 0.4 

ps4) fi 3.9 

56 9 6.2 

32 33 1.0 

42 

26 

26 

18 

13 

35 48 0 Dar 

23 46 0.5 

18 

25 14 14 

25.8 18.3 2.6 

DS 8.5 2 

14.0 19.0 yay? 

a 54 6.1 


' Based on estimated return date to capture site; some subjects may have returned sooner than indicated. 


* Radio collared subjects 


ing. If we only include bears that died of natural causes 
the mortality rate of marked bears drops to 4.5%. The 
impact of these losses on the local Black Bear popu- 
lation remains unknown. In Labrador, for much of 
the 1980s and 1990s a single Black Bear license pro- 
vided a quota of five bears per year; this has since been 
reduced to two bears per year (D. Blake personal 
communication). It is unclear how many Black Bears 
were harvested annually in the study area when the 
Black Bear quota was at its highest, but given the weak 
fur market and lack of a sport hunt, the harvest was 
probably low compared to elsewhere in Labrador. 
Estimates by provincial wildlife officials suggest 
that bear mortalities in the study area averaged about 
one per year from 1998-2003 (F. Phillips, D. Blake 
personal communication). In 2003, mining camp offi- 
cials recorded over 300 bear sightings. In total four 
Black Bears were captured, of these two were trans- 


located and two were destroyed (D. Lampe personal 
communication). While not rigorous, this informa- 
tion seems to suggest that in 2003 Black Bears were 
still relatively abundant in the study area. 

Although not an initial goal of the study, we also 
tracked the effectiveness of bear translocation and 
homing. Our results were consistent with observations 
of bear homing in other areas (Rogers 1986); most 
translocated bears (75%) eventually returned to the cap- 
ture area, generally within 1-2 weeks. 

Mahoney et al. (2001) analysed body mass for Black 
Bears from various regions of North America. The 
lowest average weight for adult females were from 
Quebec at 54 kg, and the highest from insular New- 
foundland at 101 kg; the lowest average weight for 
adult males was reported in Maine at 116 kg, and the 
highest from insular Newfoundland at 178 kg. The 
median weight of adult females from our study (48 kg) 


a 


BLACK BEAR ECOLOGY 


CHAULK, BONDRUP-NIELSEN, AND HARRINGTON 


2005 


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72 THE CANADIAN FIELD-NATURALIST Vol. 119 


TABLE 5. Visual observations (n=185) of Gross Habitat use as percent use by radio collared black bears 
(n=10) near Voisey’s Bay Labrador, from June to November, 1996. Habitats were visually classed as 
either barren, forest, or other at time of data collection. * 


Bear ID N Barren Forest Other 
(%) (%) (%) 
AMBOl1 11 DES 63.6 9.1 
AFBO2 21 40.7 51.9 7.4 
AFBO04 28 85.7 £43 0.0 
AFBO06 19 26.3 (ses, 0.0 
AFBO7! 22 68.2 31.8 0.0 
AFBO8 20 10.0 80.0 10.0 
AFBO09 1% 11.8 70.6 17.6 
SMB10 14 28.6 57.1 14.3 
SMBI11 14 304) 64.3 0.0 
SFB16 13 69.2 30.8 0.0 
Count 185 80 95 10 
Mean 18.5 40.4 53.8 5.8 
Median 18 322. 60.4 Sul 
SD 5.8 25:7 2A 5 6.8 


' AFBO7 was caring for two cubs in 1996. 


TABLE 6. Summary of Chi-square goodness of fit test for evidence of selection. The 95% CI were based on the normal 
approximation to the binomial distribution, with a Bonferroni correction for multiple testing. MCP home range was used to 
delineate habitat availability. 


Bear ID Habitat Area (obs- Proportion 95% Cl 
(Km?) | Observed Expected exp)’/exp Available Lower Upper 
AFBO02 
Barren 10.07 5 8.23 Jee 0.18 -0.00 O22 
Forest 35.97 32 29.38 0.23 0.65 0:35 0.87 
Other 9.05 8 7.39 0.23 0.16 0.04 0.31 
Total 55.09 45 45 
yo La3 
df 2 
P-value 0.46 
AFB04 
Barren 8.91 6 10.42 1.87 0.43 0.05 0.53 
Forest 8.6] 19 10.07 4.78 0.42 0.63 1303 
Other 3:01 I Sy 1.80 0.15 -0.06 0.16 
Total 20:53 24 24 
es 8.45 
df 2 
P-value 0.01 
AFBO08 
Barren 13.37 i) PS) 6.08 O31 0.02 0.23 
Forest Pa os) 48 35.63 4.29 0.64 0.75 0.97 
Other 2.43 | 3.14 1.68 0.06 -0.02 0.06 
Total 43.45 56 56 
a 12.05 
df 2 
P-value 0.00 


were slightly lower than those reported from Quebec reported for Black Bears in northern Labrador by 
by Mahoney et al. (2001). The median weight of adult = Veitch and Harrington (1996). 

males in our study (120) was slightly higher than males Daily activity patterns arise in response to seasonal 
from Maine. In general the median weight for our sub- and diurnal variation in the environment (Nielsen 
jects was comparable to the lowest average weights 1983). Black Bears are generally considered diurnal, 
reported by Mahoney et al (2001) and to weights a view substantiated by Amstrup and Beecham (1976), 


2005 


Lindzey and Meslow (1977), and Lariviere et al. (1994). 
Lariviere et al. (1994) found that Black Bears in Gas- 
pesie National Park commenced daily activity approx- 
imately 0.5 hours after sunrise, and ceased activity ap- 
proximately 2.5 hours after sunset. These results are 
similar to those observed in our study. The activity sen- 
sors revealed greatest activity in late afternoon several 
hours before sunset, and minimal activity about 2-3 
hours after sunset. However, our bears appeared to 
resume activity later in the night, several hours before 
dawn. Unfortunately the batteries in the GPS collars 
failed in mid-August and the short life span of the 
GPS collars prevented analysis across seasons. 

In Maine, Schooley et al. (1994) reported denning 
periods ranging from 134 to 197 days, with entry oc- 
curring in October and November and emergence 
occurring in April. In Alberta, Tietje and Ruff (1980) 
reported numbers that translate into a median denning 
period of 171 days, with average den entry occurring in 
October and emergence in April. In northern Labrador, 
Veitch (1994) reported denning periods ranging from 
180-220 days. The estimated denning period for bears 
in our study area ranged from 148 to 222 days (median 
= 204 days), with the median emergence occurring in 
May. Our data for both entry and emergence are limit- 
ed due to the frequency of monitoring during the fall 
and spring periods. As reported our data are similar 
to those for barren-ground Black Bears, but in reality 
they may be intermediary between bears in northern 
Labrador and elsewhere. 

In northern areas where large hollow trees are un- 
common, bears tend to use excavated dens lined with 
plant material (Fuller and Keith 1980; Tietje and Ruff 
1980; Klenner and Kroeker 1990). All known dens in 
our study were excavated, den roofs were supported 
by the root systems of the adjacent vegetation, and all 
entrances faced south or southwest, possibly to mini- 
mize exposure to north winds and to increase expo- 
sure to sunlight. 

MCP home ranges based on 1.5 months of GPS loca- 
tion data were used to determine the habitat availability 
boundary of each subject. At least two adult females 
occupied forested habitat disproportional to their avail- 
ability during the period July-August. The most com- 
monly used habitat classes appeared to be Spruce/Fir/ 
Dwarf Shrub, Birch Thicket, Black Spruce/Lichen, and 
Tuckamore. However, subject sample size (n = 3), loca- 
tion sample size (n = 24-56), time frame (1.5 months), 
and geographical extent of base mapping place restric- 
tions on generalizing habitat selection behaviors to 
other individuals. If Black Bears in the study area prefer 
forest to barren habitats it did not seem to be supported 
by visual observations of the VHF radio collared bears, 
where 3 of 10 were found more often on barrens than 
in forests. However, the visual observations occurred 
over a large region and habitat availability could not be 
determined, so statistical analysis of habitat use rela- 
tive to availability could not be conducted. 


CHAULK, BONDRUP-NIELSEN, AND HARRINGTON: BLACK BEAR ECOLOGY 


Schwartz and Franzmann (1989) found that Black 
Bears in Alaska accounted for 80% of Moose preda- 
tion and 70% of Moose mortality; however, even there 
Moose predation provided only a small proportion of 
the overall Black Bear diet. We observed two inci- 
dences of ungulate predation/scavenging in our study 
during 1996; these corroborate findings elsewhere 
(Schwartz and Franzmann 1989; Veitch and Krizan 
1996). 

At the outset of the study we were curious which 
characteristics, if any, were similar: to the barren ground 
Black Bears in northern Labrador, to bears elsewhere, 
or were intermediary between the two. We found that 
habitat use was likely to be intermediary between bears 
in northern Labrador and bears from other regions of 
North America. Cub production, homing, daily activity 
patterns, and den site construction, were similar to that 
reported from other regions of North America. How- 
ever, the small body size was most similar to barren 
ground Black Bears from more northerly regions of 
Labrador. 


Acknowledgments 

Thanks for personal communications are due to D. 
Blake and F. Phillips (Department of Forest Resources 
and Agrifoods, Newfoundland and Labrador), A. Veitch 
(Department of Resources, Wildlife and Economic 
Development, Northwest Territories), and D. Lampe 
(Labrador Inuit Association, Newfoundland and Labra- 
dor (NL). Thanks for support of this project are extend- 
ed to J. Rowell (Labrador Inuit Association, NL), B. 
Napier (INCO, Ontario), P. Trimper (Jacques Whitford 
Environment Limited, NL), and L. Cijnak-Chubbs. 
Thanks for assistance in the field are due to G. Chaulk, 
M. Michelin, B. Duffett, B. Mactavish, M. Davis and 
the many pilots at Canadian Helicopters who partici- 
pated in this study. Thanks are also extended to the 
Labrador Inuit Association and the Nasivvik Centre, 
University of Laval, for providing graduate funding. 


Documents Cited (marked * in text) 

JWEL. 1997. Ecological Land Classification. Voisey’s Bay 
Environmental Baseline Technical Data Report. Voisey’s 
Bay Nickel Co., St. John’s. xx pages. 

Kohn, B. E., and R. E., Rolley. 2000. Black Bear Population 
Analyses. Wisconsin Wildlife Surveys 10: xx pages. Avail- 
able online at http://www.wildwisconsin.com/bearpop.html. 

Lotek Engineering Inc. 1996. The GPS Animal Location 
System: Users Manual. Release 2, Newmarket, Ontario. 
51 pages. 

Rogers, L. L. 1993. The role of habitat quality in the natu- 
ral regulation of black bear populations. Proceedings of 
the 4° Western Black Bear Workshop: 95-102. Yosemite 
National Park, California. Technical Report NPS/NRWR/ 
NRTR-93/12. 


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174 


Coy, P. L., and D. L. Garshelis. 1992. Reconstructing re- 
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Dimmick, R. W., and M. R. Pelton. 1996. Criteria of sex and 
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Eason, T. H., B. H. Smith, and M. R. Pelton. 1996. Re- 
searcher variation in collection of morphometrics on Black 
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Elowe K. D., and W. E. Dodge. 1989. Factors affecting Black 
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Environment Canada. 1989. Canadian Climate Normals, 
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Environment Canada. 1998. Natural light tables for the sun: 
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62° 60' west. Moncton, New Brunswick. 

Fuller, T. K., and L. B. Keith. 1980. Summer ranges, cover 
type use and denning of Black Bears near Fort McMurray, 
Alberta. Canadian Field-Naturalist 94: 80-82. 

Jonkel, C. J., and I., Cowan. 1971. The black bear in the 
spruce-fir forest. Wildlife Monographs 27: 1-57. 

Klenner, W., and D. W. Kroeker. 1990. Denning behavior 
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Lariviere, S., J. Huot, and C. Samson. 1994. Daily activity 
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Lindzey, F. G., and E. C. Meslow. 1977. Home range and 
habitat use by Black Bears in Southwestern Washington. 
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Mahoney, S. P., J. A. Virgl, and K. Mawhinney. 2001. 
Potential mechanisms of phenotypic divergence in body 
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Moen. R., J. Pastor, T. Cohen, and C. C. Schwartz. 1996. 
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Neu, C. W., C. Randall, and J. M. Peek. 1974. A technique 
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Nielsen, E. T. 1983. Relation of behavioural activity rhythms 
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Rogers, L. 1986. Homing by radio-collared Black Bears, 
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Schooley, R. L., C. R. McLaughlin, G. J. Matula, and W. 
B. Krohn. 1994. Denning chronology of female Black 
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Schwartz, C. C., and A. W. Franzmann. 1989. Bears, wolves, 
moose and forest succession: Some management consid- 
erations on the Kenai Peninsula, Alaska. Alces 25: 1-10. 

Tietje, W. D., and R. L. Ruff. 1980. Denning behaviour of 
Black Bears in boreal forest of Alberta. Journal of Wildlife 
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1536. 

Veitch, A. M. 1994. Black bear research on the barren-grounds 
of the Northeastern Labrador peninsula, 1989-1993. Osprey 
Doe TNT. 

Veitch, A. M., and P. K. Krizan. 1996. Black bear predation 
on vertebrates in Northern Labrador. Journal of Wildlife 
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Veitch, A. M., and F. H. Harrington. 1996. Brown bears, 
black bears, and humans in northern Labrador: a historical 
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Received 18 May 2001 
Accepted 7 March 2005 


Long-tailed Weasel, Mustela frenata, Movements and Diggings 
in Alfalfa Fields Inhabited by Northern Pocket Gophers, Thomomys 
talpoides 


GILBERT PROULX 


Alpha Wildlife Research & Management Ltd., 229 Lilac Terrace, Sherwood Park, Alberta, Canada T8H 1W3; email: gproulx @ 
alphawildlife.ca 


Proulx, Gilbert. 2005. Long-tailed Weasel, Mustela frenata, movements and diggings in alfalfa fields inhabited by Northern 
Pocket Gophers, Thomomys talpoides. Canadian Field-Naturalist 119(2): 175-180. 


Little is known about the movements of Long-tailed Weasels (Mustela frenata) in alfalfa (Medicago spp.) fields inhabited by 
Northern Pocket Gophers (Thomomys talpoides). In central Alberta, I intermittently followed the movements of Long-tailed 
Weasels during two consecutive winters. Three types of movements were observed: straight-line movements across the 
fields; sinuous movements along the edges of the field; and arc movements within the field, over concentrations of Northern 
Pocket Gopher burrow systems. Arc movements were 0.6-28-m-wide at their base, and extended from 6 to 45 m into the 
field. Movements of Long-tailed Weasels into the fields were accompanied by fresh diggings in Northern Pocket Gopher 
burrow systems, which became inactive thereafter. This study suggests that Long-tailed Weasels may have a cognitive map 
of the distribution of Northern Pocket Gophers in their home range. 


Key Words: Long-tailed Weasel, Mustela frenata, Northern Pocket Gopher, Thomomys talpoides, Alfalfa fields, cognitive 


map, movements, Alberta. 


In agricultural areas, Long-tailed Weasels (Mustela 
frenata) are associated with waterways and habitats 
with abundant prey, such as fields with pocket gopher 
(Geomys spp. and Thomomys spp.) burrow systems 
that are inhabited by several small terrestrial species 
(Vaughan 1961; Gamble 1980; Whittaker et al. 1991). 
Criddle and Criddle (1925) suggested that, in winter, 
weasels entered burrow systems to kill the resident 
pocket gopher and other rodents. Andersen and Mac- 
Mahon (1981) suggested that a low survival rate in a 
population of Northern Pocket Gophers (Thomomys 
talpoides) was probably due to a sudden influx of 
weasels. Proulx and Cole (1998) identified Northern 
Pocket Gopher remains in Long-tailed Weasel scats. 
Although a predator-prey relationship between the two 
species undoubtedly exists, little is known about Long- 
tailed Weasel movements in fields inhabited by North- 
ern Pocket Gophers. 

Weasel foraging behavior has been described as a 
random search during which the animals explore every 
likely place for small prey (Powell 1978; King 1989). 
Soper (1964) considered that they wander erratically 
from place to place, visiting vegetation clumps, burrow 
openings, and boulders. In this study, I hypothesized 
that Long-tailed Weasels would investigate Northern 
Pocket Gopher burrow systems as they encounter them 
along their wanderings. 


Study Area and Methods 

This project study was carried out in two study areas 
along the Vermilion River approximately 4 km from 
Vegreville (53°N, 112°W), Alberta. Study areas were 


175 


bordered by thickets of willow (Salix spp.) and aspen 
(Populus tremuloides), high grass, and alfalfa (Medica- 
go spp.) fields inhabited by Northern Pocket Gophers. 


1998-1999 

Study Area I was surveyed for Long-tailed Weasel 
tracks 10 times from 18 November to 22 December 
1998, when snow was < 30 cm deep and without crust, 
and 15 times from 14 January to 30 March 1999, when 
snow was deeper and with crust. Air temperatures 
ranged from -29°C to 9°C. A female adult Long-tailed 
Weasel (track length: 3.8 cm; distance between jumps 
= 45 cm) was captured on 6 December in a mesh trap, 
anesthetized in a veterinary clinic with isoflurane, ear- 
tagged, and radio-collared (2.5 g collar with loop an- 
tenna; Holohill Systems, Ottawa, Ontario). Because 
weasels are very sensitive to handling and collaring 
(Delattre et al. 1985), the animal was kept in captivity 
overnight, then released at the original capture site, and 
intermittently located from 6 to 22 December, at which 
time the radio-collar ceased to emit. Radio-telemetry 
was used to locate the Long-tailed Weasel’s dens. Snow- 
tracking was used to determine the extent of the Long- 
tailed Weasel movements along the creek and in the 
alfalfa field. When following the animal, fresh dig- 
gings into the dirt were flagged and re-visited the fol- 
lowing spring. Tracks of Mink (Mustela vison) and 
Short-tailed Weasel (Mustela erminea) were also iden- 
tified on the basis of tracks and stride characteristics 
(Murie 1975; Rezendes 1992). 

Study Area II was surveyed four times from 24 No- 
vember to 22 December 1998, and six times from 15 


176 


January to 11 March 1999. The tracks of one Long- 
tailed Weasel, judged to be a male according to the size 
of its tracks (track length: 6.3 cm; distance between 
jumps = 65 cm), were found and followed. Because 
of very cold nights, often below -20°C, no attempt was 
made to live-trap the animal. Tracks of Short-tailed 
Weasels were observed across the alfalfa field. 

Animal movements and diggings were originally 
plotted on 1:20 000 scale maps drawn from air photos 
(Alberta Environmental Protection, Air Photo Service, 
Edmonton, Alberta). In April 1999, immediately after 
snow melting, the distribution of Northern Pocket 
Gophers in Study Area I was determined using dirt 
mounds (piles of soil pushed to the surface of the 
ground by pocket gophers). Because the animals’ tun- 
nel network may extend past the border delineated by 
the mounds (Proulx et al. 1995), boundaries delineated 
with dirt mounds were extended another 3 m to the 
outside. In Study Area II, Northern Pocket Gopher 
burrow systems were sparsely distributed across the 
alfalfa field and were not plotted on a map. Sites where 
Long-tailed Weasel tracks were recorded in the alfalfa 
field were visited in the spring and inspected for signs 
of Northern Pocket Gophers. 


2000 

One Long-tailed Weasel was followed four times in 
Study Area I from 4 to 31 January 2000, when tem- 
peratures ranged from -15°C to -5°C. A crust was pres- 
ent after 8 January, and tracking conditions were diffi- 
cult. Although tracks were identical to those of the 
female studied the previous year, it is not sure that this 
was the same animal. Overlap between Long-tailed 
Weasel tracks and Northern Pocket Gopher burrow 
systems was determined using the presence of mounds, 
and the pocket gopher distribution map developed 
the previous year. 


Results 
Habitats 

In 1998-1999, the female’s movements encompassed 
an 8-ha area comprising approximately 1.3 km of the 
Vermilion River, and a continuous alfalfa field (Fig- 
ure 1). Two dens were found at the interface of the 
riparian shelterbelt and the alfalfa field. The first one 
was located within the alfalfa field, 5 m from its edge, 
and it had three active openings in a Northern Pocket 
Gopher burrow system. The second den was located 
at the base of a rose (Rosa spp.) bush, in the grass- 
dominated riparian vegetation, 3 m from the edge of 
the alfalfa field. In 1999-2000, a Long-tailed Weasel 
was found within the same 8-ha area, with tracks lead- 
ing to a den, in a coarse woody debris pile at the base 
of a large willow (Figure 1). 

The male’s movements encompassed an 18-ha area 
comprising approximately 1.2 km of the Vermilion 
River, and a continuous alfalfa field (Figure 2). Tracks 
led to consecutive holes in the river’s bank, which may 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


have been its den, near a Beaver (Castor canadensis) 
dam. 


Straight-line movements 

Straight-line movements were recorded in 1998-1999 
when the female crossed from one side of the field to 
the other, at narrows (Figure 1). In 2000, a 43-m long 
straight-line movement into the field led to a digging 
site. The male crossed the field in a straight line to 
enter bush areas on either side of narrows, or to inves- 
tigate the tracks of a smaller weasel (possibly Mustela 
erminea). 


Sinuous movements 

In 1998-1999, the female traveled in a sinuous way 
along the creek and the field’s edge, investigating 
woody debris, ground openings, and boulders. Sinu- 
ous movements consisted of a series of zigzags with 
bends and turns approximately 30 cm on each side of 
an imaginary central line (Figure 3). Such movements 
were extensive along the field’s edge and extended 
from one den to the other (Figure 1). In 2000, sinuous 
movements were recorded on the south side of the 
study area. The male Long-tailed Weasel also traveled 
all along the edges of the river and the field in a sinu- 
ous manner. 


Arc movements 

In November and December 1998, the female Long- 
tailed Weasel interrupted its continuous, sinuous move- 
ments along the riparian shelterbelt-field ecotone with 
pronounced arcs that extended into the alfalfa field 
(Figure 3). These arcs began and ended at the edge of 
the field. Seven arcs of various sizes were recorded 
(Figure 1). From the center of their base to the apex, 
they extended from 6 to 45 m into the field (x = 19.6 m, 
standard error = 5 m). The width at their base ranged 
from 0.6 to 28 m (x = 10.2 + 4 m). Twenty-five North- 
ern Pocket Gopher concentrations were found in the 
alfalfa field (Figure 1). Five (71%) of the seven Long- 
tailed Weasel arc movements overlapped Northern 
Pocket Gopher concentrations. Long-tailed Weasel 
tracks circled or crossed Northern Pocket Gopher 
mounds (Figure 4) and earth plugs (holes filled up 
with soil by pocket gophers returning from the surface 
to the underground tunnel of their burrow system). 

Six male arc movements were recorded in 1998 
(Figure 2). They extended from 5 to 52 m into the field 
(x = 14.8 + 7.5 m). The width at their base ranged 
from 3 to 38 m (x = 12.5 + 5.3 m). Three (50%) of the 
arc movements overlapped Northern Pocket Gopher 
burrow systems. 

In 2000, 10 arc movements were recorded when 
tracking the Long-tailed Weasel in Study Area I (Fig- 
ure 1). They extended from 6 to 20 m into the field 
(length x : 11.2 + 2 m; width x : 6.8 + | m). The North- 
ern Pocket Gopher distribution had expanded consid- 
erably across the field since the previous year, and all 
arc movements overlapped burrow systems. 


| 
| 
| 


2005 


Field 


Field 


PROULX: LONG-TAILED WEASEL MOVEMENTS AND DIGGINGS 


177 


ie 


Field 


*. 
TeuReetag * 


ene 
N 
, 

500 


Northern Pocket Gopher concentration areas 


Long-tailed Weasel arc movements 
Long-tailed Weasel straight-line movements 


Long-tailed Weasel den 


FiGurRE |. Dens and arc movements of Long-tailed Weasels in winters 1998-1999 and 2000 in Study Area I, Vermilion 


River, Vegreville, Alberta. 


Diggings 

Twenty fresh diggings by the female Long-tailed 
Weasel were flagged during the 1998 surveys, all in the 
alfalfa field and mostly along arc movements. Seven- 
teen (85%) of them were in Northern Pocket Gopher 
burrow systems (Figure 4). The holes were still open 
when re-visited in April 1999. 


Discussion 

This study validated the hypothesis that Long-tailed 
Weasels investigated Northern Pocket Gopher burrow 
systems as they encountered them. However, encoun- 
ters were not the result of random wanderings. Long- 
tailed Weasels left their line of travel along the field 
ecotone to enter the field and investigate it with arc 
movements of various lengths and widths that over- 
lapped specific Northern Pocket Gopher concentra- 


tions. Peters (1978) and Powell (2000) suggested that 
carnivores had cognitive maps of where they live. Pow- 
ell (1978, 1994) showed that the Fisher (Martes pen- 
nanti), another mustelid, did not use the space within 
its home range randomly. Powell documented cases 
where the Fisher crossed in straight-line areas of low 
prey availability, but spent more time investigating 
Porcupine (Erethizon dorsatum) winter dens. The arc 
movements recorded in this study suggest that Long- 
tailed Weasels also have a cognitive map of the distri- 
bution of potential preys. Long-tailed Weasel apparent- 
ly directed their movements to active Northern Pocket 
Gopher burrow systems that they opened and entered. 
While Northern Pocket Gophers keep their burrow sys- 
tem closed from outside intruders (Witmer et al. 1999), 
the fact that the systems opened by the Long-tailed 
Weasels were not plugged back suggests that resident 


178 THE CANADIAN FIELD-NATURALIST Vol. 119 


e 
et, 
iat f.\ 
e 
rs 
, 
r) 
Pod 


: Field we 
B BG Peer a 500 
‘ A ee 
i m 
eae (\ Long-tailed Weasel arc movements 
Long-tailed Weasel straight-line movements 
se Long-tailed Weasel den 
o"*, River boundary 
e 
o 


FIGURE 2. Arc movements and possible den of a male Long-tailed Weasel in winter1998-1999 in Study Area I, Vermilion 
River, Vegreville, Alberta. 


Northern Pocket Gophers either abandoned their bur- 

row system or more likely were killed by Long-tailed 

Weasels. Simms (1979) suggested that Long-tailed 

Weasels in western Canada preyed heavily on North- 

ern Pocket Gophers. He estimated minimum passable 

tunnel diameter for average-sized male Long-tailed 

Weasels to be 4.3 cm, which is smaller than the mini- 

mum average diameter of 5.8 cm for tunnels of Tho- 

momys spp. (Miller 1957; Bonar 1995*). 

The sinuous movements of Long-tailed Weasels Pos 

along riparian shelterbelts and field edges are in agree- RAID: 

ment with previous reports on the meandering nature of ~ 
Long-tailed Weasel movements (Quick 1944; Wobeser \/J \/ }}.}»}»}».}..}.}. VY VG 
1966). Powell (1978) pointed out that weasel foraging 
behavior was characterized by frequent direction 
changes. Such movements would increase their chances 


to encounter small mammals, and might help avoid- —_ Figure 3. Schematic representation of Long-tailed Weasel 
ing predators (Powell 1978). sinuous and arc movements. 


Are 
movement 


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2005 


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PROULX: LONG-TAILED WEASEL MOVEMENTS AND DIGGINGS 


WE 


179 


be eo 4 
Tes 
¢ at rie # ” 


é 


FIGURE 4. Snow-covered Northern Pocket Gopher mound opened (hole diameter: 5 cm) by a female Long-tailed Weasel in 
winter 1998-1999 in Study Area I, Vermilion River, Vegreville, Alberta. 


Snowtracking is an advantageous technique as it al- 
lows one to study how Long-tailed Weasels use micro- 
environments. More snowtracking data should be gath- 
ered on Long-tailed Weasels in alfalfa fields to better 
understand their relationship with Northern Pocket 
Gophers. Unfortunately, the collection of such data is 
not an easy task. It is difficult to find alfalfa fields that 
are in proximity to water, and are inhabited by both 
species. On the other hand, a better understanding of 
the use of alfalfa fields and Northern Pocket Gopher 
populations by Long-tailed Weasels in a well-developed 
agricultural area such as central Alberta may be vital to 
ensure that this mustelid does not become threatened, 
as was feared a few decades ago (Gamble 1982*). 


Acknowledgments 

I am thankful to Alberta Conservation Association, 
Ducks Unlimited, and Red Deer River Naturalist Club 
for funding this project. I am grateful to C. Bohoychuk 
for allowing us to work on his land. I thank veterinarian 
Louis Kwantes, Park Veterinary Clinic in Sherwood 
Park, for anaesthetizing the female weasel, and Pamela 
J. Cole and Nicole A. Proulx for technical assistance. 
I thank Pauline Feldstein, Alpha Wildlife, and Roger 
A. Powell, North Carolina University, for reviewing 
an earlier version of this manuscript. This paper bene- 
fited from the comments of one anonymous reviewer 
and David Nagorsen. 


Documents Cited (marked * in text) 

Bonar, R. E. 1995. The Northern Pocket Gopher — most of 
what you thought you might want to know, but hesitated 
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Report 9524-2806 MTDC, Missoula, Montana. 

Gamble, R. L. 1982. Status report on the prairie long-tailed 
weasel Mustela frenata longicauda. COSEWIC report, 
Ottawa. 


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Gamble, R. L. 1980. The ecology and distribution of Mustela 
frenata longicauda Bonaparte and its relationships to other 
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Manitoba, Winnipeg. 

King, C. 1989. The natural history of weasels and stoats. 
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Miller, M. A. 1957. Burrows of the Sacramento Valley pocket 
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Proulx, G., M. J. Badry, P. J. Cole, R. K. Drescher, A. J. 
Kolenosky, and I. M. Pawlina. 1995. Summer above- 
ground movements of Northern Pocket Gophers, Tho- 
momys talpoides, in an alfalfa field. Canadian Field- 
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tailed Weasel, Mustela frenata longicauda, scats. Canadian 
Field-Naturalist 112: 345-346. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


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1991. Factors influencing Meadow Vole, Microtus penn- 
sylvanicus, distribution in Minnesota. Canadian Field- 
Naturalist 105: 403-405. 

Witmer, G. W., R. E. Marsh, and G. H. Matschke. 1999. 
Trapping considerations for the fossorial pocket gopher. 
Pages 131-139 in Mammal trapping. Edited by G. Proulx, 
Alpha Wildlife Research & Management Ltd., Sher- 
wood Park, Alberta. 

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Received 9 January 2004 
Accepted 14 March 2005 


Long-tailed Duck, Clangula hyemalis, Eider, Somateria spp., and Scoter, 
Melanitta spp., Distributions in Central Alaska Beaufort Sea Lagoons, 
1999-2002 


LYNNE. NogEL!?, STEPHEN R. JOHNSON?, and GILLIAN M. O’ DOHERTY! 


'LGL Alaska Research Associates, Inc., 1101 East 76" Avenue, Suite B, Anchorage, Alaska 99518 USA 
*LGL Limited, 9768 2" Street, Sidney, British Columbia V8L 3Y8 Canada 
Present addresses. ENTRIX, Inc., 1600 A Street, Suite 304, Anchorage, Alaska 99501 USA; e-mail: Inoel @entrix.com 


Noel, Lynn E., Stephen R. Johnson, and Gillian M. O’ Doherty. 2005. Long-tailed Duck, Clangula hyemalis, eider, Somateria 
spp., and scoter, Melanitta spp., distributions in Central Alaska Beaufort Sea Lagoons, 1999-2002. Canadian Field- 
Naturalist 119(2): 181-185. 


During July and August 1999-2002, distributions of Long-tailed Ducks (Clangula hyemalis), eiders (Somateria spp.) and 
scoters (Melanitta spp.) were documented in three barrier island-lagoon systems in the central Alaska Beaufort Sea. Concentration 
areas for each species were determined during 16 aerial surveys. Kernel density procedures were used to delineate 75% and 
50% “activity” or concentration areas for all three species. Long-tailed Ducks were 13 times more numerous than eiders and 
38 times more numerous than scoters. The Long-tailed Duck 75% activity area encompassed all three lagoon systems and was 
three times as large as the eider activity area and one-third larger than the scoter activity area. Eider activity areas were located 
only in the eastern lagoon, and scoter activity areas were located only in the western lagoon. Density contours showed patterns 
of repeated habitat use for sea ducks over the four years of sampling and improve our understanding of sea duck habitat use 


within Beaufort Sea barrier island-lagoon habitats. 


Key Words: Clangula hyemalis, Melanitta, Somateria, molt, normal kernel density, Arctic, barrier island, Alaska. 


Up to 30 000 molting waterfowl aggregate during 
mid-July through early September along the mainland 
and barrier island shorelines in the central Beaufort 
Sea, Alaska (Figure 1; Johnson et al. 2005). Since 1990, 
Long-tailed Ducks (Clangula hyemalis), the predom- 
inant waterfowl species in the central Alaska Beaufort 
Sea lagoon systems (Johnson and Gazey 1992), have 
shown a declining trend in total numbers during the 
male molt period in these lagoons (i.e., mid July to late 
August) (Fischer et al. 2002; Johnson et al. 2005). 
During 1999-2002, distributions of waterfowl in the 
barrier island-lagoon systems between Spy Island and 
Flaxman Island, Alaska, were documented (Figure 1). 
Normal kernel density contours, a smoothed density 
estimation function (Powell 2000), for the combined 
distributions are presented to delineate “activity” or 
concentration areas for Long-tailed Ducks, eiders and 
scoters within these lagoons. 


Methods 
Aerial Surveys 

During each survey, the western lagoon, Stefansson 
Sound, and eastern lagoon subdivisions of the study 
area were surveyed on the same day (Figure 1; Johnson 
1990; Johnson and Gazey 1992). Lagoon systems were 
surveyed in varying orders, but all transects within a 
lagoon system were surveyed before moving to the 
next lagoon system, and the entire survey area was 
flown within a 5-hour period. A total of 16 surveys 
totaling 59 hours was conducted over four years. Four 


surveys each were flown during 30 July to 26 August 
1999 and | to 24 August 2000, three surveys were flown 
during 23 July to 11 August in 2001, and five surveys 
were flown during 20 July to 14 August 2002. 

Aerial surveys were conducted in a single-engine 
fixed high-wing aircraft (Cessna 206) on floats. The 
survey crew consisted of a pilot and two observers. 
One observer sat in the right front seat and the other in 
the left rear seat. Survey altitude was approximately 
45 m above ground level and ground speed was ap- 
proximately 180 km/h. Transect width was 400 m total, 
200 m on each side of the aircraft. 

The surveys were during the peak of the male Long- 
tailed Duck molt (flightless) period from 15 July to 
21 August (Johnson and Richardson 1981; Johnson 
1985; Johnson and Gazey 1992). Because Long-tailed 
Ducks are known to concentrate along the barrier is- 
lands in the late afternoon and evening (Johnson and 
Richardson 1981; Johnson 1982, 1985), surveys were 
scheduled as late in the day as practical, generally 
from 14:00 to 19:00. Surveys on days with high winds 
(>35 km/h) were postponed until winds, wave height, 
and chop diminished, thereby improving the observers’ 
ability to see birds on the water (Johnson 1990; John- 
son and Gazey 1992). 


Survey Data 

Tape recorders were used to record all bird obser- 
vations. Continuous audio-tape recordings included 
information on transects, observations, survey weather 


181 


182 


SPY 
“ SIMPSON LAGOON 
arc ne J 


—< = WESTERN 
~~. LAGOON 


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Po ~ S\ f / 


Wis 


BAY. 


THE CANADIAN FIELD-NATURALIST 


Nae S%p 
| PRUDHOE “24, 


Vol. 119 


Transect Line 
> ES 


\ 


FiGureE |. Location of aerial survey transects used to record Long-tailed Duck, eider and scoter distribution and abundance 
in the barrier island-lagoon systems between Spy Island and Flaxman Island, Alaska, July and August 1999-2002. 


conditions, and 30-second intervals (time periods) 
marked by a timer. A notebook computer with map- 
ping software linked to a Global Positioning System 
(GPS) receiver recorded the flight line at one-second 
intervals during surveys. Observers synchronized their 
watches with GPS satellite time, and these times (re- 
corded for transects and time periods) were used to 
geo-reference survey data. 


Normal Kernel Density Contours 

Density contours were computed and mapped using 
the ArcView® (ESRI, 1996, Redlands, California) Ani- 
mal Movement software extension (Hooge and Eichen- 
laub 1997*; Silverman 1986; Worton 1989; Seaman 
and Powell 1991). “Activity” or concentration areas 
were represented by density contours delineating 75% 
and 50% of birds (Powell 2000). Computations were 
based on the total number of on-transect individuals 
summarized for each 30-second time-period over the 
entire survey area. 

Density contours were calculated for Long-tailed 
Ducks, all eiders and all scoters. Most eiders (99% of 
eiders identified to species) were Common Eiders 
(Somateria mollissima v-nigrum) with very small num- 
bers of King Eiders (Somateria spectabilis). Most 
scoters (98% of scoters identified to species) were Surf 


Scoters (Melanitta perspicillata) with small numbers 
of Black Scoters (Melanitta nigra) and White-winged 
Scoters (Melanitta fusca). 


Results 

Long-tailed Ducks were 13 times more numerous 
than eiders and 38 times more numerous than scoters 
(Table 1). The 75% activity area for Long-tailed Ducks 
included parts of survey areas in all three lagoon sys- 
tems and was three times as large as the eider activity 
area and one-third larger than the scoter activity area 
(Figure 2, Table 1). The only eider activity area was 
in the eastern lagoon, and the only scoter activity area 
was in the western lagoon (Figure 2). The 50% con- 
tour area was about one-third to three-quarters of the 
size of the 75% contour area depending on the species 
(Figure 2, Table 1). 

The eider activity area was used mainly by large 
flocks of molting males, and smaller groups (<10 birds) 
of resting females and females with broods aggregated 
along barrier-island or mainland shorelines. Scoters 
occurred in scattered mixed feeding flocks with Long- 
tailed Ducks in mid-lagoon habitats in the western 
lagoon. Eider and scoter activity areas did not overlap 
(Figure 2). 


TABLE |. Activity area size (km7), total bird density (number per km?) and total number of Long-tailed Ducks, eiders, and 
scoters during July and August 1999-2002 in the barrier island-lagoon systems between Spy Island and Flaxman Island, 


Alaska. 
Total birds 75% activity area 
16 surveys km? 
Long-tailed Ducks 181224 642 
Eiders 13708 225 
Scoters 


Density 50% activity area Density 
Number per km? km? Number per km? 
243 238 388 

38 97 54 


4762 423 


2005 NOEL, JOHNSON, AND O’ DOHERTY: LONG-TAILED DUCK, EIDER, AND SCOTER IN ALASKA 183 


LAGOON. Long-tailed Ducks 


~ 


STEFANSSON 75% 50% 


10 20 
kilometers 


EASTERN 
LAGOON 


. 7 
Yj tof LZ RI 


75% 50% 


kilometers 


EASTERN 
LAGOON 


75% 


10 20 
kilometers 


EASTERN 


FIGURE 2. Long-tailed Duck, eider and scoter activity areas for 75% and 50% of total birds during July and August 
1999-2002 in the barrier island-lagoon systems between Spy Island and Flaxman Island, Alaska. 


184 


Discussion 

Long-tailed Duck distributions in barrier island- 
lagoon systems reflected proximity to both lagoon for- 
aging habitats and resting habitats along the barrier 
island and mainland shorelines (Johnson 1990; John- 
son and Gazey 1992; Johnson et al. 2005). Eiders and 
scoters both were more limited in their distributions 
than Long-tailed Ducks (Figure 2). Eider activity area 
was limited to the eastern lagoon, while scoter activity 
area was within the western lagoon (Figure 2). Eider 
distributions reflected foraging, nesting and resting 
habitat use, but scoter distributions reflected only for- 
aging and resting habitat use. Only the Long-tailed Duck 
75% activity area included the Stefansson Sound sub- 
division of the study area. 

Long-tailed Ducks are less specific in their prey 
selection than are either eiders or scoters (Nilsson 
1972; Stott and Olson 1973, Sanger and Jones 1984). 
Long-tailed Duck prey may be either nektonic or 
benthic (Johnson 1984; Stott and Olson 1973). In the 
Simpson Lagoon area, preferred prey are crustaceans 
and bivalves in 2—3 m water depths, where prey densi- 
ties are highest (Johnson 1984). Given the less specific 
nature in their foraging, and observations of mixed- 
species foraging flocks of Surf Scoters and Long-tailed 
Ducks in the western lagoon, competition and mutual 
exclusion between Long-tailed Ducks and scoters is 
unlikely. 

Eiders are primarily benthic feeders and, although 
they may be more specific in their food habits than 
Long-tailed Ducks, they are more general than scot- 
ers (McGilvrey 1967; Stott and Olson 1973; Cantin 
et al. 1974; Vermeer 1981). The eider activity area in 
the eastern lagoon encompassed the deeper waters (up 
to 6 m), with crustacean and bivalve prey (Woodward- 
Clyde Consultants 1996*) known to be preferred by 
this species (Nilsson 1972). 

Aggregations of eiders are often associated with 
breeding habitats (Savard et al. 1999). Common Eider 
nesting activities in our study area are concentrated 
on the Stockton Islands in the eastern lagoon system 
and on Egg and Stump islands in the western lagoon 
system (Figure 1; Johnson 2000). The 50% eider activ- 
ity contour in our study included the Stockton Island 
group; however, the center of this activity area was 
directly associated with the larger foraging flocks of 
eiders, not with known nesting areas (Figure 2). 

There was no overlap in activity areas for eiders and 
scoters (Figure 2). Although some of the same foods 
are preferred by these two species, different foraging 
habitats were selected. The lack of overlap between 
eider and scoter activity areas may reflect differential 
habitat use similar to that described elsewhere (Savard 
et al. 1999). Scoter distributions are known to be asso- 
ciated with foraging habitats rich in bivalves (Nilsson 
1972; Vermeer 1981). Although bivalves are known 
to occur throughout the barrier island-lagoon systems 
in the Alaska Beaufort Sea, fine-scale prey sampling 
across all known feeding habitats has not yet occurred. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Direct links between sea duck distribution and prey 
availability in Beaufort Sea barrier island-lagoon hab- 
itats have only been documented in a few studies 
(Johnson and Richardson 1981; Johnson 1984). Nev- 
ertheless, this study and others (Johnson et al. 2005) 
show strong patterns of repeated use of discrete lagoon 
habitats by sea ducks. These patterns are likely linked 
to food availability during the mid-summer molt period. 
This information will be helpful when designing sea 
duck mitigation and monitoring programs as devel- 
opment occurs in barrier island-lagoon habitats in the 
Alaska Beaufort Sea. 


Acknowledgments 

Preparation of this manuscript and field studies 
were funded by BP Exploration (Alaska) Inc. (BP) and 
the Point Thomson Unit Owners (BP, ExxonMobil 
Production Company, Conoco Phillips Alaska, Inc. and 
others). W. Streever, D. Trudgen, A. Erickson, and W. 
Cullor, BP Environmental Studies Group, provided 
suggestions concerning this study and assistance with 
logistical arrangements. W. Streever, BP Environmen- 
tal Studies Group Leader, and G. Robilliard, ENTRIX, 
Inc. provided comments that improved this manuscript. 
J. Helmericks piloted the survey aircraft. In addition 
to the authors, T. Olson, I. Helmericks, R. Rodrigues, 
and M. Bentley served as observers. 


Documents Cited (marked * in text) 

Hooge, P. N., and B. Eichenlaub. 1997. Animal movement 
extension to ArcView. Version 1.1. Alaska Biological Sci- 
ence Center, U.S. Geological Survey, Anchorage, Alaska. 

Seaman, D. E., and R. A. Powell. 1991. Kernel Home Range 
Estimation Program. Version 4.26. United States Geolog- 
ical Survey, Biological Resources Division, Port Angeles, 
Washington. 

Woodward-Clyde Consultants. 1996. The 1995 Northstar 
Unit sampling program. Unpublished report prepared for 
BP Exploration (Alaska) Inc., Anchorage, Alaska by 
Woodward-Clyde Consultants, Anchorage, Alaska. 


Literature Cited 

Cantin, M., J. Bédard, and H. Milne. 1974. The food and 
feeding of common eiders in the St. Lawrence estuary in 
summer. Canadian Journal of Zoology 52: 319-334. 

Fischer, J. B., T. J. Tiplady, and W. W. Larned. 2002. 
Monitoring Beaufort Sea waterfowl and marine birds, 
aerial survey component. U.S. Fish and Wildlife Service, 
Division of Migratory Bird Management, Anchorage, 
Alaska. OCS Study MMS 2002-002. Available from Na- 
tional Technical Information Service, NTIS Order Number: 
PB2002-102377. 

Johnson, S. R. 1982. Continuing investigations of long- 
tailed ducks (Clangula hyemalis L.) during the molt peri- 
od in the Alaskan Beaufort Sea. Pages 547-563 in Envi- 
ronmental Assessment of Alaskan Continental Shelf, Final 
Report of Principal Investigators. Volume 23. Bureau of 
Land Management and National Oceanic and Atmospheric 
Administration, Outer Continental Shelf Environmental 
Assessment Program, Boulder, Colorado. Available from 
National Technical Information Service, NTIS Order Num- 
ber: PB85-212595/AS. 


2005 


Johnson, S. R. 1984. Prey selection by Oldsquaws in a Beau- 
fort Sea lagoon, Alaska. Pages 12-19 in Marine birds: their 
feeding ecology and commercial fisheries relationships. 
Canadian Wildlife Service Special Publication, Ottawa, 
Ontario. 

Johnson, S. R. 1985. Adaptations of the long-tailed duck 
(Clangula hyemalis L.) during the period of molt in Arctic 
Alaska. International Ornithological Congress 18: 530-540. 

Johnson, S. R. 1990. Monitoring Beaufort Sea waterfowl 
and marine birds. Report MMS 90-0048 by LGL Alaska 
Research Associates, Inc., Anchorage, Alaska, for Minerals 
Management Service, Anchorage, Alaska. Available from 
National Technical Information Service, NTIS Order Num- 
ber: PB91-235242. 

Johnson, S. R. 2000. Pacific Eider. Pages 259-275 in The nat- 
ural history of an arctic oil field. Edited by J. C. Truett and 
S. R. Johnson. Academic Press, San Diego, California. 

Johnson, S. R., and W. Gazey. 1992. Design and testing of 
a monitoring program for Beaufort Sea waterfowl and 
marine birds. Report MMS 92-0060 by LGL Limited, 
Sidney, British Columbia, for Minerals Management Ser- 
vice, Anchorage, Alaska. Available from National Technical 
Information Service, NTIS Order Number: PB93-135903. 

Johnson, S. R., and W. J. Richardson. 1981. Barrier island 
lagoon ecological process studies: Final Report, Simpson 
Lagoon. Part 3, Birds. Pages 109-383 in Environmental 
Assessment of the Alaskan Continental Shelf, Final Reports 
of Principal Investigators. Volume 7. Biological Studies. 
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Bureau of Land Management and National Oceanic and 
Atmospheric Administration, Outer Continental Shelf 
Environmental Assessment Program, Boulder, Colorado. 
Available from National Technical Information Service, 
NTIS Order Number: PB82-192113/AS. 

Johnson, S. R., L. E. Noel, W. J. Gazey, and V. C. Hawkes. 
2005. Aerial monitoring of marine waterfowl in the central 
Alaskan Beaufort Sea. Environmental Monitoring and 
Assessment 108: 1-43. 


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185 


McGilvrey, F. B. 1967. Food habits of sea ducks from the 
north-eastern United States. Wildfowl Trust Annual Report 
18: 142-145. 

Nilsson, L. 1972. Habitat selection, food choice, and feeding 
habits of diving ducks in coastal waters of south Sweden 
during the non-breeding season. Ornis Scandinavica 3: 
55-78. 

Powell, R. A. 2000. Animal home ranges and territories and 
home range statistics. Pages 65-110 in Research techniques 
in animal ecology: Controversies and consequences. Edited 
by L. Boitani and T. K. Fuller. Columbia University Press, 
New York, New York. 

Sanger, G. A., and R. D. Jones, Jr. 1984. Winter feeding 
ecology and trophic relationships of Oldsquaws and White- 
winged Scoters on Kachemak Bay, Alaska. Pages 20-28 in 
Marine birds: their feeding ecology and commercial fish- 
eries relationships. Canadian Wildlife Service Special Pub- 
lication, Ottawa, Ontario. 

Savard, J.-P. L., J. Bedard, and A. Nadeau. 1999. Spring 
and early summer distribution of scoters and eiders in the 
St. Lawrence River estuary. Pages 60-65 in Behavior and 
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sen, and G. J. Robertson. Canadian Wildlife Service Occa- 
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Silverman, B. W. 1986. Density estimation for statistics and 
data analysis. Chapman and Hall, London, England. 

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Received 26 May 2004 
Accepted 11 March 2005 


Winter Habitat Use by Moose, Alces alces, in Central Interior British 
Columbia 


GILBERT PROULX! and RHONDA M. Kariz2 


'Alpha Wildlife Research & Management Ltd., 229 Lilac Terrace, Sherwood Park, Alberta T8H 1W3 Canada 
Driftwood Forestry Consulting, 16880 Schefer Drive, Prince George, British Columbia V2K 5L4 Canada 


Proulx, Gilbert, and Rhonda M. Kariz. 2005. Winter habitat use by Moose, Alces alces, in central interior British Columbia. 
Canadian Field-Naturalist 119(2): 186-191. 


In central British Columbia, recent epidemics of Mountain Pine Beetle (Dendroctonus ponderosae) have resulted in the use 
of expansive clearcut areas to remove infested mature and old Lodgepole Pine (Pinus contorta) stands. This study aimed to 
determine if Moose (Alces alces) use late-successional Lodgepole Pine stands in mid- to late-winter. Moose activity and 
habitat use was determined during February-March track surveys in 2000 (60 km) and 2001 (55.7 km). In 2000 (69 tracks) 
and 2001 (313 tracks), Moose track distribution differed significantly (P < 0.05) from random. They were significantly more 
abundant than predicted in young stands (dominated by Picea spp.), or early seral stages; they were less abundant than pre- 
dicted in mature and old Lodgepole Pine stands. It is unlikely that harvesting late-successional Lodgepole Pine stands would 


affect Moose winter habitat supply. 


Key Words: Alces alces, Moose, snowtracking, Lodgepole Pine, Pinus contorta, British Columbia. 


Most work suggests that a proper interspersion of 
food and shelter in winter can be an important and pos- 
sibly limiting feature of Moose (Alces alces) habitat 
(Welsh et al. 1980; Proulx and Joyal 1981; Peek et al. 
1987). Timber harvesting may have a positive effect 
on Moose by creating early seral stages rich in browse. 
On the other hand, it may be detrimental to Moose, 
particularly in winter, if a mosaic of varied cover types 
and regeneration sites is not properly maintained 
(Telfer 1970). 

In central British Columbia, recent epidemics of 
Mountain Pine Beetle (Dendroctonus ponderosae) 
have resulted in the infestation of at least 4.2 million 
hectares of mature and old Lodgepole Pine (Pinus con- 
torta) stands (generally > 80 years), and the use of ex- 
pansive clearcut silviculture programs to recover the 
timber (Readshaw 2003*). Moose is a species highly 
valued by local residents and native groups, and the 
B.C. Ministries of Forests and Water, Lands, and Air 
Protection voiced their concerns regarding the removal 
of Lodgepole Pine cover and the possible reduction 
in Moose winter habitat. 

This study aimed to determine if Moose used 
late-successional Lodgepole Pine stands in mid- to 
late-winter, when relatively deep snow and low temper- 
atures force animals to seek canopy cover with well- 
developed understories (Thompson and Stewart 1997). 
Because late-successional pine stands do not usually 
offer a multi-storied cover and a well-developed under- 
story, we hypothesized that Moose would make little 
use of late-successional lodgepole pine. 


Study Area 
The study area was located near Burns Lake (54°14'N, 
125°46'W), in central British Columbia, 350 km west 


186 


of Prince George. It encompassed an 80 km? area that 
is part of the Sub-Boreal Spruce Biogeoclimatic (SBS) 
Zone (Meidinger and Pojar 1991). The climate of the 
SBS zone is continental, and is characterized by sea- 
sonal extremes of temperatures, i.e., severe winters and 
relatively warm, moist, and short summers (Meidinger 
et al. 1991). The SBS is part of the Canadian Boreal 
Forest Region (Krajina 1965). Upland coniferous for- 
ests dominate the sub-boreal landscape. Hybrid White 
Spruce (Picea engelmanii x glauca) and subalpine fir 
(Abies lasiocarpa) are the dominant climax tree spe- 
cies. Lodgepole Pine is common in mature forests in 
the drier parts of the zone, and both Lodgepole Pine 
and Trembling Aspen (Populus tremuloides) pioneer 
the extensive early-seral stands (Meidinger et al. 1991). 
Wetlands are common and dot the landscape in poorly 
drained, postglacial depressions or river ox-bows. Wet- 
land community types include sedge (Carex spp.) 
marshes, and fens with birch (Betula spp.), willows 
(Salix spp.), Black Spruce (Picea mariana), and hybrid 
spruce (P. glauca x P. mariana) (Meidinger et al. 1991). 

Winter was generally harsher in 2000 than in 2001. 
In February, in situ data collections indicated that tem- 
peratures ranged from -28 to 2°C in 2000. They ranged 
from -24 to 9°C in 2001, but most days were between 
-4 and 0°C. During both years, surface conditions con- 
sisted of loose granular snow without crust, and hooves 
of Moose penetrated deeply into the snow. Mean snow 
depths of most habitats were significantly greater in 
2000 than in 2001; on average, mean snow accumu- 
lation was 11.7 cm deeper in 2000 than in 2001 (Proulx 
and Kariz 2001*). In February 2000, mean snow depths 
approximated 43 cm in coniferous forests and 62 cm 
in openings/immature stands. In 2001, they averaged 
33 and 52 cm in the same respective habitats. 


2005 


Methods 
Track surveys 

Moose activity and habitat use was determined 
during two consecutive surveys each year: from 4-11 
February and 24 February-1 March, 2000, and 30 Jan- 
uary-5 February and 1-6 March, 2001. Each survey 
consisted of seven line transects crossing the study 
area. The same transects were used during both years, 
with some deviations due to open water in 2001. Tran- 
sect lengths also varied within and between years due 
to accessibility (e.g., extensive, recent blowdowns 
interfered with researchers’ movements) and environ- 
mental conditions (e.g., sudden change of weather with 
heavy snowfall); they ranged from 3.6 to 5.4 km, and 
were oriented to avoid paralleling roads (Cairns and 
Telfer 1980). They were plotted on forestry maps, and 
starting points were tied by compass bearings and 
distance to distinctive topographic features. Transects 
were snowshoed using a compass and 1:20 000 for- 
estry maps; linear distance along a survey transect was 
determined with a hip chain and recorded each time 
there was a change of habitat type. Because the study 
focused on assessing the use of pine stands by Moose, 
habitat classification was kept simple, and limited to 
four types based on the British Columbia Resources 
Inventory Committee’s (1998*) classes (Table 1). 

Because Moose may take advantage of packed 
snowshoe trails in their daily movements (E. Telfer, 
Canadian Wildlife Service, personal communication), 
different transects were used from one survey period to 
the other during a same year. Each year, transects of the 
second survey period were placed in between those of 
the first survey. In winter, Moose usually move <1 km 
during their daily wanderings (Phillips et al. 1973; 
Hundertmark 1997) and, in order to avoid problems 
associated with pseudoreplication (Hurlbert 1984), 
transects of a same survey period were located at least 
1 km apart. Therefore, tracks encountered in different 
transects during a survey period were likely from dif- 
ferent animals. Along transects, because it was not pos- 
sible to consistently determine if crossings were made 
by the same animal, all crossings were tallied (Raphael 
and Henry 1990*). Only fresh tracks were recorded. 


Data analyses 

All track transects were surveyed within one month, 
under similar weather conditions. Transects of the first 
and second surveys within a same year were pooled 
together for data analyses. Student t-test was used to 
compare mean numbers of tracks/transect between 
years. Pair-sampled f-testing was used to compare dif- 
ferences in the number of tracks recorded along a same 
transect from one year to the other (Zar 1999). Propor- 
tions of habitat classes traversed by survey transects 
were used to determine the expected frequency of track 
intersects per habitat class if tracks were distributed 
randomly with respect to habitat classes (Parker et al. 
1981). Chi-square statistics with Yates correction (Zar 
1999) were used to compare observed to expected fre- 


PROULX AND KARIZ: WINTER HABITAT USE BY MOOSE 


187 


quencies of track intersects per habitat class. When 
chi-square analyses suggested an overall significant 
difference between the distribution of observed and 
expected frequencies, comparisons of observed to 
expected frequencies for each habitat class were con- 
ducted using a G-test for correlated proportions (Sokal 
and Rohlf 1981). Probability values < 0.05 were con- 
sidered statistically significant. 


Results 
Transects’ vegetation composition 

A total of 60 and 55.7 km were snowshoed in 2000 
and 2001, respectively. Vegetation composition along 
transects was similar during both years (x7: 0.85, df: 
5) P= 0105) (Figure 1), 


Frequency of Moose tracks/transect by year 

On average, the number of Moose tracks/transect 
was significantly lower (t = 4.79, P < 0.05) in 2000 
(n: 14 transects, x = 4.9 tracks, standard deviation = 
4.2 tracks) than in 2001 (n: 14, x = 22.3 + 12.9 tracks). 
The number of tracks/transect ranged from 0 to 17 in 
2000, and from 4 to 46 in 2001. Differences observed 
in the number of tracks recorded along a same transect 
from one year to the other were highly significant (¢ = 
5.16; P< O01). 


Frequency of Moose tracks per habitat type 

In 2000, 69 fresh Moose tracks were encountered. 
The observed frequency of tracks per habitat class was 
significantly different (y? = 23.62, df: 5, P = 0.001; 
Figure 2) from a random distribution of tracks among 
habitat classes. Tracks were significantly more abun- 
dant than predicted in young stands (G = 4.19, df: 1, 
P <0.05) (Figure 2). They were less abundant than pre- 
dicted in late-successional pine stands (G = 8.21, df: 
1, P < 0.01) (Figute'2). 

In 2001, 313 fresh Moose tracks were encountered. 
The observed frequency of tracks per habitat class was 
significantly different (y’= 81.06, df: 5, P < 0.001; 
Figure 3) from a random distribution of tracks among 
habitat classes. Tracks were significantly more abun- 
dant than predicted in openings and immature | stands 
(G = 7.83, df: 1, P < 0.01), and in immature 2, scrub 
and pole stands (G = 3.85, df: 1, P< 0.05). They were 
less abundant than predicted in late-successional pine 
stands (G = 35.12, df: 1, P< 0.001) (Figure 3). 


Discussion 

During the cold winter of 2000 and the warmer win- 
ter of 2001, this study validated our expectation that 
Moose make little use of late-successional Lodgepole 
Pine in mid- to late-winter. The low use of late-succes- 
sional pine stands may be explained by a generally 
poor understory with few deciduous shrubs for browse, 
and the lack of a multi-storied vertical cover (Raphael 
et al. 1992*). 

During the two winters, Moose used a mosaic of 
habitat patches that offered deciduous shrubs for food 
and coniferous canopy for shelter. However, habitat 


188 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |. Habitat types used in the study of Moose habitat use in February-March 2000 and 2001 in central British Colum- 
bia (after British Columbia Resources Inventory Committee’s (1998) classes). 


Forest type 


Deciduous 


Coniferous 
= Pure 
— Mixed 


Coniferous-deciduous 
Habitat type 


Openings and immature | stands 


Immature 2, pole, and scrub 


Young 


Late successional 


Characteristics 


Crown closure = 10%, deciduous species > 75% 


Crown closure = 10%, coniferous species > 75% 

When = 80% of the coniferous cover is provided by one species. 

When the coniferous cover is provided by more than one species, neither 
species = 80% 


Crown closure = 10%, neither type > 75% 
Description 


0-10 years old. Open areas with sparse shrubs, and replanted clearcuts with 
trees < 2 m high. 


11-40 years old. Immature 2 stands represent new forests following a natural 
or anthropogenic disturbance, with trees = 2 m high. Pole stands are thick 
stands of trees (7.5 to 12.4 cm diameter at breast height), usually with little 
understory. Trees compete with one another and other plants for light, water, 
nutrients, and space to the point where most other vegetation and many trees 
become suppressed and die. Scrubs are typical lowlands and bogs with short 
Black Spruce and/or willow or alder thickets. 


40-80 year-old forests consisting mainly of spruce-dominated stands. 
Achievement of dominance by some trees and death of other trees leads to 
reduced competition that allows understory plants to become established. The 
forest canopy has begun differentiation into distinct layers. Vigorous growth 
and a more open and multi-storied stand than in the pole stage. 

= 81 years old forests consisting mostly of mature stands with even canopy of 
trees, with or without coarse woody debris down and leaning logs. A few old 
stands with tall and large canopy trees, canopy gaps, large snags, large downed 
woody debris, and developed understories. A second cycle of shade tolerant 
trees may have become established. Multi-layered canopy and developed 
understories usually missing in late-successional pine stands. 


Openings & Immature2, Young Late- Late- Late- 

immature 1 pole, and successional successional successional 

scrub pine spruce and _ coniferous- 

mixed deciduous 
coniferous and 

; deciduous 

Habitat types 
FIGURE |. Distance traveled through each habitat type during Moose track surveys in central 


British Columbia, in February-March 2000 and 2001. 


2005 PROULX AND KARIZ: WINTER HABITAT USE BY MOOSE 189 


25 


C Observed 
@ Expected 


Number of Moose tracks 


Openings & Immature2, Young Late- Late- Late- 
immature 1 ___ pole, and successional successional successional 
scrub pine spruce and _coniferous- 
mixed deciduous 
coniferous and 
Habitat types deciduous 


FIGURE 2. Observed and expected number of Moose tracks per habitat type in central British 
Columbia, February-March 2000. 


100 
90 L] Observed 
@ Expected 
80 
DH 
3 70 
5 
» 60 
= 
Ss 30 
ei 40 
vo 
2 
E 30 
Z 
20 
10 
0 Pees mates 
Openings Immature2, Young Late- Late- Late- 
immature! pole, and successional successional successional 
scrub pine spruce and  coniferous- 
mixed deciduous 
coniferous and 
Habitat types deciduous 


FIGURE 3. Observed and expected number of Moose tracks per habitat type in central British 
Columbia, February-March 2001. 


mosaics used by Moose varied in composition be- stands. Moose apparently sought habitats that offered 
tween years. Moose were less active in 2000 than in both cover and food. The use of such habitats may 
2001, as suggested by the markedly lower number of — occur with relatively shallow snow (Sandegren et al. 
recorded tracks. In February-March 2000, Moose 1985; Hundertmark 1997) and low temperatures 
tracks were found in a variety of young and mature (Schwartz and Renecker 1997), and has been reported 


190 


in many regions (Stevens 1970; Eastman 1974; Rolley 
and Keith 1980). In February-March 2001, Moose 
tracks were more abundant than predicted on the basis 
of a random distribution in openings with vegetation, 
immature stands and scrub. Moose can experience 
heat stress in winter when temperatures rise above 
—5°C (Schwartz and Renecker 1997); they may have 
used these habitats because they are rich in browse, 
but are also more open, windier, and cooler than forests 
(Renecker and Schwartz 1997). 

It is unlikely that harvesting late-successional 
lodgepole pine stands to address Mountain Pine Bee- 
tle infestations would impact on Moose winter habi- 
tat supply. However, Moose may use pine stands to 
travel between wetlands and young forests, and cross 
clearcuts (Proulx, personal observation). Then, if pine 
stands are retained as connectivity corridors across the 
landscape, non-clearcut silviculture treatments may be 
warranted to sanitize beetle-infested stands and pro- 
vide traveling Moose with protective cover. 


Acknowledgments 

We thank the British Columbia Ministries of En- 
vironment, and Water, Land and Air Protection for 
inquiring about Moose winter habitat, and Canadian 
Forest Products Ltd. (Canfor), Houston Operations, for 
funding the study. We thank Carl Vandermark, Canfor, 
for his interest in this study, and Joel Nicholson for 
technical help. We thank two anonymous referees for 
their valuable comments. 


Documents Cited (marked * in text) 

British Columbia Resources Inventory Committee. 1998. 
Standard for terrestrial ecosystem mapping in British 
Columbia. Resource Inventory Committee, Victoria. 

Proulx, G., and R. Kariz. 2001. Moose winter habitat in 
Canfor’s Operating Area in Lakes Forest District, British 
Columbia, 1999-2001. Alpha Wildlife Research & Man- 
agement Ltd. report prepared for Canadian Forest Prod- 
ucts Ltd., Houston, British Columbia. 

Raphael M. G., and S. E. Henry. 1990. Preliminary sugges- 
tions for monitoring Marten in the Rocky Mountains 
region. United States Department of Agriculture, Forest 
Service. Pacific Northwest Forest Science Laboratory, 
Olympia, Washignton. 6 pages. 

Raphael, M., S. L. Marquardt, and R. H. Barrett. 1992. 
Evaluating stand conditions to support integrated sylvicul- 
tural pescriptions for timber and wildlife: snags and old 
growth. Pages 1-20 in Proceedings habitat futures work- 
shop, Edited by J. B. Nyberg and W. B. Kessler, Pacific 
Experimental Forest, British Columbia Mininstry of For- 
ests, Victoria. 

Readshaw, K. 2003. Timber supply analysis, Mountain Pine 
Beetle infestation. British Columbia Ministry of Forests, 
Victoria. Mimeograph. 


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movement patterns and range use in northwestern Min- 
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Seasonal Moose migrations related to snow in Sweden. 
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the North American Moose. Edited by A. W. Franzmann 
and C. C. Schwartz, Wildlife Management Institute. Book, 
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37-46. 

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Thompson I. D., and R. W. Stewart. 1997. Management of forest harvesting. Proceedings North American Moose 
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D.C. 


Welsh, D. A., K. P Morrison, K. Oswald, and E.R. Thomas. Received 24 November 2003 
1980. Winter habitat utilization by Moose in relation to Accepted 14 March 2005 


Facts from Faeces: Prey Remains in Wolf, Canis lupus, Faeces 
Revise Occurrence Records for Mammals of British Columbia’s 
Coastal Archipelago | 


MICHAEL H. H. Price!, Curis T. DARIMONT!*, NEVILLE N. WINCHESTER!, and PAUL C. PAQUET2? 


‘Department of Biology, University of Victoria. P.O. Box 3020, Victoria, British Columbia, V8W 3N5 Canada 

*Raincoast Conservation Society. P.O. Box 26, Bella Bella, British Columbia VOT 1B0 Canada 

Faculty of Environmental Design, University of Calgary, 2500 University Drive Northwest, Calgary, Alberta T2N 1N4 
Canada 


Price, Michael H. H., Chris T. Darimont, Neville N. Winchester, and Paul C. Paquet. 2005. Facts from faeces: prey remains 
in Wolf, Canis lupus, faeces revise occurrence records for mammals of British Columbia’s coastal archipelago. 
Canadian Field-Naturalist 119(2): 192-196. 


Archipelagos often harbour taxa that are endemic and vulnerable to disturbance. Conservation planning and research for 
these areas depend fundamentally on accurate and current taxonomic inventories. Although basic ecological information is 
in its infancy, the temperate rainforest islands of coastal British Columbia are undergoing rapid human-caused modification, 
particularly logging. We report herein new mammal records for these islands as determined by prey remains in the faeces of 
Wolves (Canis lupus), the area’s apex mammalian terrestrial predator. Of particular interest is our detection of Marten 
(Martes americana) on islands previously inventoried and island occupancy by Moose (Alces alces), which have apparently 
migrated recently to coastal British Columbia. Remains in faeces provided valuable new species occurrence information, 
but more extensive and focused inventories are required to generate predictions of island occupancy by mammals based on 
biotic and abiotic landscape features. 


Key Words: Wolves, Canis lupus, conservation, distribution, faeces, inventories, islands, mammals, archipelago, British 


Columbia. 


Oceanic archipelagos often harbour taxa that are en- 
demic, highly vulnerable to disturbance, and prone to 
extinction (Burkey 1995; Alcover et al. 1998). Sensible 
land-use planning often is constrained by a paucity of 
data on the distribution and abundance of organisms 
(Soulé and Kohm 1989), which may be particularly 
severe on islands. Biotic inventories provide critical 
information for conservation planning but frequently 
are not conducted because of time, financial, or other 
constraints (Kremen 1994). Due to accelerating rates 
of habitat loss, species inventories may be the founda- 
tion by which to measure the extent of human influ- 
ences on extinction-prone biotas (Cook et al. 2001). 
Conversely, in poorly studied systems facing imminent 
disturbance, inventories may instigate timely conser- 
vation strategies and identify important areas for con- 
servation research. 

The numerous islands constituting British Colum- 
bia’s coastal archipelago are nearly pristine yet face 
increasing pressure from development, particularly log- 
ging (Darimont and Paquet 2000*, 2002). Although 
earlier distribution reports have been valuable (McCabe 
and Cowen 1945; Cowen and Guiguet 1975; Craig 
1990*, Nagorsen 1990), a paucity of fundamental eco- 
logical information still remains (such as complete 
mammal community records) for British Columbia’s 
islands. In contrast, mammalian distribution on the 
adjacent Alexander Archipelago of Southeast Alaska 
has been well described (MacDonald and Cook 1996) 
and notable patterns of biogeography (e.g., Conroy et 


al 1999) and endemism (e.g., Cook and MacDonald 
2001; Fleming and Cook 2002; Small et al. 2003) 
have emerged. 

Accounts of mammal occurrence on islands are not 
always systematic but often dynamic, reflecting iterative 
coalescence of various collection records, accounts, 
and personal observations (e.g., MacDonald and Cook 
1996). We extend this process here by benefiting from 
the foraging habits of coastal British Columbia’s apex 
mammalian predator, the Wolf (Canis lupus), which has 
a wide potential niche. 

During summers 2000 and 2001, we collected Wolf 
faeces along British Columbia’s coast between the 
Kshwan Valley (55 °37'N, 129 °48'W) in the north and 
the Koeye River (51°46'N, 127°53'W) in the south 
(Figure 1). Our study area and sampling procedures 
are well described elsewhere (Darimont and Paquet 
2000*, 2002; Darimont et al. 2004). Herein we iden- 
tify mammalian prey occurring in faeces collected on 
islands only. We compared hair in faeces with vouch- 
er samples and also used dichotomous keys (Mathiak 
1938; Mayer 1952; Stains 1958; Price 2003*). We 
compared these records of prey remains on islands 
with existing information regarding mammal occur- 
rence for the British Columbia archipelago (McCabe 
and Cowen 1945; Cowen and Guiguet 1975; Craig 
1990*, Nagorsen 1990) and herein report differences. 

Wolves may deposit faeces from prey items con- 
sumed on other landmasses, but we consider this poten- 
tial bias negligible. Although not well described, esti- 


[eZ 


2005 PRICE, DARIMONT, WINCHESTER, AND PAQUET: FACTS FROM FAECES 193 


Islands Sampled 


Pearse 
Wales 
Dundas 
Dunira 
Stephens 
Smith 
Lewis 

. McCauley 
. Anger 

. Banks 

. Pitt 

. Hawkesbury 
. Farrant 

. Gribbell 

. Fin 

. Dewdney 

. Campania 
. Gil 

. Princes Royal 
. Moore 

. Aristazabal 
. Pooley 

. Roderick 

. Chatfield 


E. 
2. 
a 
4. 
5. 
6. 
F. 
8 
=, 
10 


_ Cunningham Waglisla 


oi) 4 
. Campbell (Bella Bella) 
. Denny # Lary 


. Goose Group 
. Hunter 


130 


Southeast 
Alaska 


Vancouver 


Island vi ia 


Victoria 


OF” 


° ° 
128 126w 


FiGurE 1. Study area and islands surveyed for mammal occurrence in British Columbia’s coastal archipelago during summers 


2000 and 2001. 


mates of typical “gastric emptying times” in wild and 
Captive wolves are rapid, ranging from 8 to 56 hours 
(Floyd et al. 1978; Weaver 1993; Kreeger et al. 1997; 
Peterson and Cuicci 2003). Weaver (1993) noted an 
average of four defecations/day in wolves fed ad libi- 


tum. Moreover, wolves typically rest after feeding and 
limit travel for several days thereafter (Mech 1966; 
Peterson 1977; Hayes et al. 2000; Peterson and Cuicci 
2003). Finally, 65% of mammals we identified on each 
island were found in two or more scats. 


194 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |. Mammal species identified in Wolf (Canis lupus) faeces collected on 29 islands in British Columbia’s coastal 
archipelago during 2000 and 2001. Shown only are mammals not previously recorded in McCabe and Cowen (1945), Cowen 
and Guiguet (1975); Craig (1990*), or Nagorsen (1990). Species are as follows: Mule Deer (Odocoileus hemionus sitkensis); 
Pine Marten (Martes americana); Mink (Mustela vison); River Otter (Lontra canadensis); Weasel (Mustela erminea); Black 
Bear (Ursus americanus), Moose (Alces alces),; Beaver (Castor canadensis). Sampling site codes match those in Figure 1. 
Islands previously inventoried for mammal occurrence indicated by asterisk. 


Island Area 
(km?) Deer Marten 
1. Pearse 226 xX xX 
2. Wales oF Xx 
3. Dundas 160 Xx 
4. Dunira 93 
5. Stephens 78 
6. Smith aE xX 
7. Lewis 7 xX 
8. McCauley* oA, 
9. Anger 51 x 
10. Banks* 1024 x 
fi: Pace* 1349 
12. Hawkesbury 322 xX xX 
13. Farrant 50 x x 
14. Gribbell 207 x 
15. Fin 13 xX 
16. Dewdney of) x 
17. Campania* 157 
18. Gil 238 xX xX 
19. Princess Royal* 2295 Xx 
20. Moore* 5 
21. Aristazabal* 451 xX 
22. Pooley 162 x x 
23. Roderick 239 Xx 
24. Chatfield* 48 
25. Cunningham 115 x 
26. Campbell 145 
27. Denny | DA xX x xX 
28. Goose* 24 xX 
29. Hunter* 399 


Early distribution reports (above) collectively inven- 
toried 10 of 29 islands surveyed in this study. Of partic- 
ular note are our new accounts for Pine Marten (Martes 
americana) on Banks, Princess Royal, and Aristazabal 
Island (Figure 1), which previous studies did not detect. 
MacDonald and Cook (1996) reported (non-introduced) 
occupancy of Marten on nine Southeast Alaskan is- 
lands, adjacent to British Columbia’s coast. Conroy 
et al. (1999) commented on their absence there from 
relatively small islands (< 232 km’). In British Colum- 
bia, we identified Marten on four islands smaller than 
232 km/?, including one as small as 50 km? (Table 1). 
These differences, however, may reflect island isolation 
(1.e., distance from other landmasses). Conroy et al. 
(1999) found that isolation (not size) best predicts 
species occurrence in coastal Alaska. Finally, the dis- 
tributions of animals that are trapped commercially, 
such as Marten, may also reflect introductions and har- 
vest pressure. 

Also notable is the presence of Moose (Alces alces) 
on Pitt Island. Moose have not been reported previously 


Mink 


Mammals Detected 


R. Otter Weasel B. Bear Moose’ Beaver 
4 X 
xX 
Xx > 4 4 
xX 
xX 
x xX 
xX 
X 
XxX X 
xX xX X 
x xX 


on islands of British Columbia and are thought to range 
only in localised areas of major mainland rivers and the 
heads of inlets (Nagorsen 1990; Shackleton 1999; Blood 
2000; but see Darimont et al. 2005). Mammals docu- 
mented on islands not previously surveyed included 
other representatives of Cervidae and Mustelidae, as 
well as Ursidae and Castoridae (Table 1). 

Data from faeces present potential biases and op- 
portunity. One limitation, and a function of our limited 
sampling and the dietary niche of Wolves, is a lack of 
data on other taxa (e.g., Fisher, Martes pennanti), and 
especially smaller rodents. The Deer Mouse (Pero- 
myscus maniculatus), for example, was absent from 
our samples. This species occurs on coastal islands 
and has been the subject of pioneering work on insu- 
larity in the region (McCabe and Cowen 1945), and 
continues to be a model organism elsewhere (e.g., Vuce- 
tich et al. 2001). Our “facts from faeces” approach, 
however, increases the breadth of valuable informa- 
tion available from analysing “waste” from animals 
(Putman 1984; Kohn and Wayne 1997). 


2005 


New mammalian records for coastal British Colum- 
bia have scientific value and utility in conservation de- 
bate and land-use planning. As data accumulate, we 
hope to present a coherent explanation as to what biotic 
and abiotic conditions predict mammalian diversity on 
British Columbia’s islands. Moreover, we agree whole- 
heartedly with MacDonald and Cook (1996) that, “the 
most distressing issue regarding our lack of knowl- 
edge for this vast area [coastal temperate rainforests | 
is that planners and policy makers are generating 
management decisions with great uncertainty. This is 
especially disconcerting given the accelerating rate of 
human-induced change occurring within the region’s 
ecological systems”. 


Acknowledgments 

This study took place in the Traditional Territories 
of several First Nation groups, from whom we sought 
permission before research began. We are extremely 
grateful to the Raincoast Conservation Society for 
financial and logistical support, field colleagues too 
numerous to acknowledge adequately, and skippers 
Stephen Anstee, Brian Falconer, Jean-Marc Leguerri- 
er, and Dave Lutz. The McCaw Foundation, National 
Geographic Society, Vancouver Foundation, Wilbur- 
force, World Wildlife Fund Canada, and private donors 
kindly provided funding. While preparing the manu- 
script, CTD was supported by a Natural Sciences and 
Engineering Research Council (NSERC) — Industrial 
Post-graduate Scholarship. 


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Darimont, C. T., and P. C. Paquet. 2000. The gray wolves 
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nical Report prepared for the Raincoast Conservation 
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Price, M. H. H. 2003. Foraging ecology of wolves in British 
Columbia’s coastal archipelago: the biogeography of pre- 
dation. BSc. honours thesis. University of Victoria. Victoria, 
British Columbia. 52 pages. 


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Received 12 March 2004 
Accepted 14 March 2005 


Attempted Predation of a Child by a Gray Wolf, Canis lupus, 


near Icy Bay, Alaska 


MarkK E. McNay! and Puitie W. MOONEY? 


‘Alaska Department of Fish and Game, 1300 College Road, Fairbanks, Alaska 99701-1599, USA 
?Alaska Department of Fish and Game, 304 Lake Street, Sitka, Alaska 99835-7563, USA 


McNay, Mark E. and Philip W. Mooney. 2005. Attempted predation of a child by a Gray Wolf, Canis lupus, near Icy Bay, 


Alaska. Canadian Field-Naturalist 119(2): 197-201. 


On 26 April 2000 a six-year-old boy was attacked and repeatedly bitten by a Gray Wolf (Canis lupus) in a logging camp 
near Icy Bay, Alaska. The animal’s behavior during the attack clearly contained elements of predation. The wolf was killed short- 
ly after the attack and found to be in normal physical condition; tests for rabies and canine distemper were negative. Low 
densities of ungulate prey and increased energetic demands associated with denning may have influenced the wolf’s behav- 
ior, but we believe the wolf’s habituation to people was a more significant factor contributing to the attack. Food-condition- 
ing may have facilitated the habituation process, but there was no evidence the attack resulted from a food-conditioned 


approach response. 


Key Words: Gray Wolf, Canis lupus, food conditioning, habituation, predation, wolf aggression, wolf attack, Alaska. 


Documented aggression by wild Gray Wolves (Canis 
lupus) toward humans in North America is rare and 
often characterized by bluff charges or brief attacks 
involving a single bite (Munthe and Hutchison 1978; 
Jenness 1985; Scott et al. 1985; Mech 1998; Strick- 
land 1999; McNay 2002a, 2002b [Case 15, page 12]). 
However, in April 2000 near Icy Bay, Alaska, a wolf 
pursued and attacked a six-year-old boy, then attempt- 
ed to carry the boy to nearby cover. The wolf inflicted 
19 laceration and puncture wounds on the boy’s back, 
legs, and buttocks before being driven away and killed. 
Among recent published accounts of wolf aggression 
toward humans in North America, none describe such 
persistent, aggressive behavior by a healthy wolf. 

The Alaska Department of Public Safety (ADPS) 
conducted an investigation immediately following the 
attack. Later, we conducted a separate review of the 
incident. Details of the attack were obtained from our 
interviews with five people who either witnessed the 
attack or observed the wolf’s behavior prior to the 
attack, and from the ADPS (2000*) report that includ- 
ed interviews with six additional people. We describe 
this uncommon incidence of wolf aggression and dis- 
cuss possible motivations for the attack. 


The Setting 

The attack occurred at a logging camp near Icy 
Bay, Alaska (59°58'N, 141°39'W). The camp was in 
a 13 ha clearing surrounded by dense forest. It served 
80 seasonal workers and seven permanent families. An 
unpaved road from a log sort yard on the shore of Icy 
Bay ran through the camp and continued westward. 
Gray Wolf habitat near Icy Bay is confined to about 
100 km? of Sitka Spruce/Western Hemlock (Picea sit- 
chensis/Tsuga heterophylla) forest surrounded by steep 
mountains, ice fields, and the Gulf of Alaska. A nar- 


row corridor of forest along the beach leads to more 
suitable wolf habitat 50 km to the west (Figure 1). 
The diet of Gray Wolves near Icy Bay probably con- 
tains a wide variety of foods. Moose (Alces alces) are 
migratory and no more than 20 are present at any time 
of the year. Mountain Goats (Oreamnos americanus) 
are found in steep terrain but are not often killed by 
wolves. Snowshoe Hares (Lepus americanus), Beaver 
(Castor canadensis), small rodents, and migratory birds 
are potential prey. Marine foods (fish, invertebrates, 
and marine mammals) are available as carrion on the 
beaches and Coho Salmon (Oncorhynchus kisutch) 
migrate into spawning creeks during late summer. 


The Attack 

About 9:00 a.m. on 26 April 2000, John Stenglein 
(age 6) and Keith Thompson (age 9) were playing in 
the forest edge on the camp’s north perimeter. They 
heard a noise, looked up and saw a wolf standing 3 m 
away behind a low tree branch. The wolf showed its 
teeth and growled as it crouched under the branch 
toward them. The boys remained still until the wolf 
stepped forward and snarled again. John, wearing 
oversized boots, ran slowly and awkwardly. Keith ran 
ahead calling for help. Keith’s dog, a male Labrador 
Retriever, was 50-100 m from the boys, but ran toward 
the wolf as the boys emerged from the forest (Figure 2). 

John ran about 40 m across open gravel, then stum- 
bled and fell. The dog briefly fought the wolf near the 
tree line, but the wolf disengaged and attacked John 
while the boy was on the ground. John struggled to 
escape, but the wolf lifted him off the ground, turned, 
and attempted to carry the boy toward the forest. 
Because of the boy’s size (27 kg) and loose clothing, 
the wolf had difficulty carrying the boy and it began 
dragging John toward the tree line. Four adults res- 


197 


198 


\ 50-kilometer beach corridor 
to other suitable habitat 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Icy Cape 


Gulf of Alaska 


N 


+ 


6 Kilometers 


FiGuRE |. Detail showing location of the Icy Bay logging camp relative to available wolf habitat (shaded area) in the vicinity 


of Icy Cape. 


ponded to the boys’ cries for help and approached to 
within | m while shouting and throwing rocks. The dog 
bit at the wolf’s hind legs, but the wolf focused on the 
boy and largely ignored the harassment by both the 
dog and rescuers. At one point the wolf released its 
grip on the boy after being struck by a rock, but then 
attacked again. Eventually, when the dog positioned 
itself between the child and the wolf, a rescuer grabbed 
the boy and carried him away (Alaska Department of 
Public Safety, 2000*; S. Norberg and T. Thompson, 
Icy Bay, Alaska, personal communication). 

The dog and the three remaining rescuers drove 
the wolf into the forest, but the wolf was defiant and 
reluctant to leave. Less than 10 minutes later, Keith’s 
father arrived with a rifle. He walked into the forest 
and briefly called with a predator call. Almost imme- 
diately, the wolf stepped onto a trail 80 m behind him. 
The man turned and fired once, killing the wolf (Alas- 
ka Department of Public Safety 2000*). 

A post-mortem examination revealed no apparent 
physical disabilities that would have contributed to 
the wolf’s behavior. The wolf’s size (approximately 
35 kg) appeared small compared to other wolves in 
that area, but body fat levels were normal. The stom- 
ach contained remnants of natural foods, including 


cartilage and hair, possibly from a Beaver. The wolf 
tested negative for rabies and canine distemper. The 
wolf wore a tightly fitting radio collar that had caused 
hair loss on its neck, but no abrasion to the skin 
(Blake 2000*). 


The Wolf’s History 

The wolf had been captured and fitted with a radio 
collar in March 1996 at 10 months of age (Alaska 
Department of Public Safety 2000*). It dispersed from 
its natal range (100 km west of Icy Bay) during win- 
ter 1997-1998. The radio was not heard again, but a 
radiocollared wolf, believed to be the same wolf, was 
seen twice by Alaska Department of Fish and Game 
(ADF&G) biologists in June 1998, 50 km west of Icy 
Bay. During one of those sightings the collared wolf 
was with another, uncollared wolf. 

Almost a year later, in April 1999, the collared wolf 
was seen on a logging road 18 km west of Icy Bay. The 
previous day, Mooney witnessed a truck driver throw 
food to an uncollared female wolf at the same site. 
Following that feeding incident, the logging company 
reemphasized their policy prohibiting the feeding of 
wildlife and there were no further known cases of 
wolves being fed. 


2005 


McNay AND MOoNEY: ATTEMPTED PREDATION BY A WOLF 


199 


“wolf shot 


| and 


kille 


initial 
1 encounter 


74 


4 


= —| wolf attack 


al 


FIGURE 2. Aerial photo of the Icy Bay logging camp showing location of the wolf encounter and attack sites. Photo by P. Mooney. 


Over the next year, the collared wolf exhibited in- 
creasingly bold behavior. It was seen at least seven times 
by camp residents along the north perimeter of camp, 
on the road, and near the log sort yard. Between Sep- 
tember 1999 and April 2000, people encountered the 
wolf at close range at least four times, but it never ap- 
proached or demonstrated aggressive behavior toward 
people. However, the wolf was suspected in an attack 
that seriously injured a dog in summer 1999. 

The pattern of sightings suggests the wolf occupied 
a territory and was not a transient. At 5 years of age 
most wolves have passed through a dispersal period 
during which they disassociate from their natal pack 
and travel alone before finding a mate (Fuller et al. 
2003). The collared wolf’s dispersal began at the nor- 
mal age of 2 years. The sightings of the collared wolf 
and a female wolf at the same location on consecutive 
days in April 1999 suggests a mated pair, and on 2 July 


2000 a female wolf with five pups crossed the road 
6 km from the camp in front of an approaching vehicle. 
One of the pups was photographed by the driver. That 
sighting confirmed a den or rendezvous site was near- 
by. Those pups would have been born in April or early 
May 2000, about the time of the attack. 

The collared wolf was often seen near the sort yard 
so it routinely traveled the 15 km between the denning 
area and the sort yard. A dry streambed along the north- 
ern perimeter of the camp provided a natural travel 
route. Residents frequently saw the collared wolf near 
camp in early summer, but not in late summer of 1999. 
That change in activity coincided with increased food 
availability as salmon moved into streams 6 km from 
the camp. 

The camp’s garbage was collected and incinerated 
daily. The Alaska Department of Public Safety inves- 
tigator inspected all garbage disposal facilities and 


200 


specifically asked 11 camp residents if they knew of 
wolves being fed or obtaining food near the camp. The 
investigator concluded there was no evidence that the 
animal was drawn into the area by active or passive 
feeding within the year prior to the attack (Alaska 
Department of Public Safety 2000*). 


Discussion 

Until the day of the attack, the collared wolf had 
never approached or acted aggressively toward people, 
but it had demonstrated increasingly fearless behavior. 
The habituation process was probably facilitated by 
the camp’s central location in the small, isolated area 
of wolf habitat. The wolf probably encountered people 
frequently, but people would have been unaware of 
the wolf’s presence during most encounters, because 
the camp, road, and sort yard were all surrounded by 
dense forest. The presence of dogs may have encour- 
aged the wolf to periodically patrol and scent mark 
along the camp’s perimeter. Company policy made it 
difficult for camp residents to hunt or trap near the 
camp or near worksites, thereby creating a defacto wolf 
protection zone where wolves were not conditioned to 
avoid humans. That pattern of frequent, low intensity 
(1.e., passive and inconsequential) encounters, irregu- 
larly spaced over a long period, is the ideal recipe for 
habituation (Kimmel 1973). 

It seems unlikely that agonistic behavior toward the 
dog precipitated the wolf’s attack because the dog was 
indoors until just before the attack and was 50-100 m 
from the boys during the wolf’s initial approach. The 
dog skirmished with the wolf at least three times, but 
suffered no injuries. During the attack, the wolf focused 
on the boy and ignored the dog, but both boys had 
been with the dog earlier that morning. It’s possible the 
dog’s scent on their clothing played a role in the wolf’s 
initial approach. 

Snarling behavior is generally associated with ago- 
nism in wolf social interactions and can precede either 
offensive or defensive aggression (Zimen 1982). High- 
ranking wolves harass subordinates by crouching and 
threatening to spring, baring their teeth, or with an open 
mouth (Mech 1970). However, the snarling behavior 
witnessed by the boys prior to the Icy Bay attack does 
not rule out predation as a motivation. Among several 
predatory attacks upon children in India, wolves stalked 
toward children then snarled or showed their teeth be- 
fore attacking (K. Rajpurohit, Wildlife Institute of India, 
personal communication). 

Predation involves a sequence of connected behav- 
iors that include: (a) orientation toward the prey; (b) 
following (i.e., stalking or rushing approach); (c) catch- 
ing, and in the case of small prey; (d) carrying (Fox 
1971). Several characteristics of the John Stenglein 
attack suggest the wolf developed a highly aroused 
predation response because (1) the wolf ventured from 
the forest to pursue the boy across an open gravel pad; 
(2) the wolf attempted to carry and drag the boy back 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


to dense cover; (3) the wolf sustained the attack in the 
presence of rescuers; (4) the wolf reluctantly left the 
attack site after the boy had been rescued; and (5) the 
wolf remained hidden in the forest near the attack site 
and responded to a predator call within minutes after 
the incident. 

When faced with conflicting stimuli, animals can 
exhibit behaviors that are compounds of separate acts 
that may be partially inhibited, ambivalent, or alternat- 
ing (Colgan 1989). The sequence and timing of events 
in the Icy Bay attack suggest the wolf hesitated before 
chasing the boy, possibly undergoing a transition from 
an agonistic threat to a predatory response. The hesi- 
tation could also reflect the lack of a prey image for 
humans, but wolves have been known to change their 
behavior toward people from submissive to dominant 
or from habituated to predaceous, based on only one 
or a few instances where they observe vulnerability 
(Strickland 1999; McNay 2002a). John ran approxi- 
mately 40 m before the wolf attacked. The boy ran 
slowly and awkwardly in oversized boots, eventually 
falling. That visual image of vulnerability, accompa- 
nied by the boy’s cries for help, was probably signifi- 
cant in eliciting the wolf’s subsequent response, which 
was no longer inhibited. The wolf’s attempt to carry 
and drag the boy away from rescuers cannot be ex- 
plained as an agonistic act and despite the possible 
agonism reflected in the wolf’s initial approach, the 
final result clearly contained elements of predation. 

In a study of wolves 150 km west of Icy Cape, wolf 
pack territories ranged from 193 to 597 km? and aver- 
aged 428 km? (J. Carnes, University of Idaho, personal 
communication), but only a 100 km? of useable habitat 
was available near Icy Bay. The sighting of a female 
wolf with pups on the road 6 km from the logging 
camp suggests the attacking wolf was paired with a 
female that produced pups, it would be unlikely another 
adult male simultaneously occupied such a restricted 
territory. 

Typically male wolves carry food to the denning 
female (Harrington and Mech 1982; Mech et al. 1999). 
That provisioning places increased energetic demands 
on the male which could affect predation rates and 
motivation. For example, predation rates by Coyotes 
(Canis latrans) on domestic sheep declined when pups 
of depredating adults were destroyed (Till and Knowl- 
ton 1983). Carbyn (1989) suggested nutritional stress 
on Coyotes during the reproductive season, in combi- 
nation with habituation to humans, contributed to 
predatory attacks by Coyotes on children. Coyotes in 
those attacks behaved similar to the Icy Bay wolf by 
repeatedly biting the victims, attempting to drag victims 
away from the attack site, and exhibiting reluctance to 
abandon their attack even when rescuers intervened. 

Some biologists who commented on the attack dur- 
ing the course of the ADPS investigation discounted 
the idea of a predatory motivation because the wolf 
was capable of quickly killing the boy, but did not 


2005 


(Alaska Department of Public Safety 2000*). However, 
interruption of the predation sequence can result in 
confusion, conflicted response, and failure to kill the 
prey (Zimen 1978). Furthermore, immediate killing 
is not a prerequisite for successful predation. In Bihar, 
India, 92 children were preyed upon by wolves be- 
tween 1989 and 1995. Wolves grabbed children by 
the neck, waist, head, chest or thigh then carried them 
away from the attack site. Twenty children were res- 
cued alive after bystanders chased the wolves (Raj- 
purohit 1999); a 4-year-old boy was rescued after vil- 
lagers pursued the wolf for 2 km (Jhala and Sharma 
1997). Even with natural prey, wolves commonly do 
not immediately kill. During Caribou (Rangifer tar- 
andus) calf mortality studies in Alaska, biologists ob- 
served wolves carry live calves to cache or feeding 
sites before killing the calf (R. Boertje, ADF&G, per- 
sonal communication). Therefore, predatory intent can- 
not be ruled out in the Icy Bay case simply because 
the boy was not killed. 

What seems most clear is that the Icy Bay wolf 
became habituated to the presence of people. Habitu- 
ation was a factor common to predatory attacks by 
Coyotes on children in North America (Carbyn 1989) 
and presumably to wolf predation on children in India 
where wolves continually live among high densities of 
people and natural foods are often scarce (Jhala and 
Sharma 1997; Rajpurohit 1999). In the Icy Bay case, 
food conditioning may have initially facilitated the habit- 
uation process, but there was no evidence the attack 
resulted from a recently reinforced, food-conditioned 
approach response. 


Acknowledgments 

We are indebted to the people of Icy Bay who 
openly discussed their experiences related to the attack. 
Those residents included Betsy Beeks, Samantha 
Norberg, Carol Simpson, Diane Stenglein, and Teresa 
Thompson. John Carnes provided unpublished data 
on wolf home range sizes. We discussed the attack 
with Trooper Marc Cloward who provided additional 
insight into findings not presented in his investigative 
report. We thank Shelley Szepanksi, Beth Lenart, and 
Laura McCarthy of ADF&G for preparation of fig- 
ures and technical editing. Steve Peterson and Patrick 
Valkenburg of ADF&G commented on an early draft 
of this paper. 


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Received 4 September 2001 
Accepted 14 April 2005 


Post-Reproductive Pacific Salmon, Oncorhynchus spp., as a Major 
Nutrient Source for Large Aggregations of Gulls, Larus spp. 


K. S. CHRISTIE and T. E. REIMCHEN 
Biology Department, University of Victoria, Victoria, British Columbia V8W 3N5 Canada; e-mail: reimchen @uvic.ca 


Christie, K. S., and T. E. Reimchen. 2005. Post-reproductive Pacific salmon, Oncorhynchus spp., as a major nutrient source 
for large aggregations of gulls, Larus spp. Canadian Field-Naturalist 119(2): 202-207. 


On the Pacific coast of North America, the most abundant vertebrate visitors to estuaries and rivers during salmon migration are 
gulls, yet the utilization of salmon nutrients by these scavengers, and subsequent ecological impacts are not well documented. 
On two forested watersheds on the central coast of British Columbia, we tracked gull abundance during the spawning period 
for two consecutive years, and estimated consumption of post-reproductive salmon carcasses and eggs, as well as guano 
production. At Clatse River, gulls (Larus glaucescens, L. argentatus, L. thayerii, L. californicus, L. canus, L. philadelphia) con- 
sumed 13-26% of total salmon carcass biomass and 29-36% of all salmon eggs deposited in the system. At Neekas River, 
gulls consumed 11-19% of salmon carcass biomass and 7-18% of total salmon eggs. Local guano production over the 60-day 
period ranged from 600 kg to 1190 kg at Clatse and from 1200 kg to 2100 kg at Neekas River, and was distributed to marine, 
estuarine, freshwater and riparian habitats. The large aggregations of gulls and subsequent nutrient cycling observed on our 
study watersheds may represent a once widespread phenomenon that is now largely reduced due to recent declines in salmon 


populations. 


Key Words: Gulls, Larus, nutrients, Oncorhynchus, salmon, spawning, consumption, eggs, carcasses, British Columbia. 


In the north Pacific, large runs of spawning salmon 
(Oncorhynchus spp.) contribute substantial quantities 
of nutrients to aquatic and terrestrial food webs (Bilby 
et al. 1996; Willson et al. 1998; Willson 2004). Nutri- 
ents from salmon carcasses are used extensively by 
many wildlife species such as bears, marten, wolves, 
eagles, gulls, and ravens and become incorporated 
into terrestrial vegetation and invertebrate communi- 
ties (Reimchen 1994*, 2000; Ben-David et al. 1998: 
Cederholm et al. 2000*; Helfield and Naiman 2001; 
Darimont and Reimchen 2002; Hocking and Reimchen 
2002). The most numerous, yet least well-studied ver- 
tebrates that feed on post-reproductive salmon are 
gulls, which congregate in the thousands on streams 
throughout the north Pacific during their southward 
autumn migration (Mossman 1958; Campbell et al. 
1990; Skagen et al. 1991). Migration and feather molt, 
both energetically demanding activities, require rapid 
accumulation of lipids (Jenni and Jenni-Eiermann 1998; 
Stocker and Weihs 1998; Hamer et al. 2002). These 
metabolic demands in addition to harsh weather condi- 
tions in the fall and winter can lead to high mortality in 
gulls, especially for juveniles (Burger 1993; Verbeek 
1993; Hamer et al. 2002). 

In this manuscript, we quantify gull abundance and 
foraging activity on two salmon streams of coastal 
British Columbia. We examine temporal shifts in abun- 
dance of gulls on each salmon stream, salmon and 
egg consumption by each species of gull and the recy- 
cling of salmon nutrients via guano production. 


Methods 
Our study was conducted on the on the central coast 
of British Columbia at the Clatse River (52°20.6'N; 


127°50.3'W) and the Neekas River (52°28.4'N; 128° 
8.0'W), both of which support spawning populations of 
Chum (Oncorhynchus keta) and Pink (O. gorbuscha) 
salmon that spawn from late August until early No- 
vember. Approximately | km from the mouth of the 
Clatse River and 2.1 km from the mouth of the Neekas 
River, 5-10 meter waterfalls act as barriers to further 
upstream migration of the salmon. These localities, 
both of which support more than 20 000 spawning 
salmon, are described in detail elsewhere (Hocking 
and Reimchen 2002; Mathewson et al. 2003). 

We made a total of 33 separate gull surveys com- 
prising 8-9 per year at each watershed throughout the 
salmon spawning period. All surveys were made during 
low tide and included both estuary and river habitats. 
Gulls were counted and identified. Large gulls were 
grouped to facilitate identification from a distance and 
later identified to species in sub-sets. Among gulls that 
were foraging, we recorded feeding technique, and ob- 
tained classifications (surface-seizing, surface-plunging, 
carcass-scavenging) from Ashmole (1971). We record- 
ed food item (carcass or eggs) consumed by sub-sets 
of foraging Mew Gulls. The study period extended 
from 9 September to 17 October in 2002 and from 25 
August to 21 October in 2003. 

Daily consumption of salmon carcasses and eggs 
was calculated for each gull species. We used the 
consumption model modified from Bishop and Green 
(2001) as follows: 


C = FMR/MEC x P x M 


where C = consumption (g day"'), FMR = field meta- 
bolic rate (KJ day'), MEC = metabolizable energy 


202 


2005 


coefficient of salmon or eggs; P = proportion of 
salmon or eggs in diet; M = mass of salmon or eggs 
(g) needed to produce | KJ energy. FMR was calcu- 
lated using the allometric equation for free-living 
seabirds from Birt-Friesen et al (1989): 


FMR = 1] Q3-08+0.667logM) 


where average body mass (M, in kg) was obtained 
from Dunning (1993) (Table 1). MEC was assumed to 
be 0.75 for both salmon flesh and eggs (Castro et al. 
1989; Bishop and Green 2001). Energy density of sene- 
scent salmon flesh is 2.95 KJ g"' (wet mass) (Hendry 
and Berg 1999), and for salmon eggs it is 7.60 KJ g'! 
(wet mass) (Jonsson et al. 1998). Calculated values of 
FMR and consumption are shown in Table 1. Total con- 
sumption by gulls per day was derived from the mean 
gull count per day for each watershed. Based on the 
surveys at each watershed, we estimated mean daily 
gull abundance using two methods (1) the mean abun- 
dance of gulls derived from the original 8-9 surveys 
and (2) the interpolated mean, where each day was as- 
signed a value based on temporal shifts in abundance. 
Although gulls occupy salmon streams throughout most 
of the 3-month spawning period, we estimated con- 
sumption for a 60-day period, the interval over which 
we had detailed data. 

We calculated the proportion of total salmon bio- 
mass consumed by gulls using total consumption esti- 
mates relative to number of salmon returning to the 
river to spawn (escapement). Salmon escapement was 
obtained for our study streams in 2002 and 2003 from 
the Department of Fisheries and Oceans (Terry Pal- 
frey, personal communication, Table 2). Average intact 
carcass mass for Pink and Chum salmon at our study 


CHRISTIE AND REIMCHEN: PACIFIC SALMON AS NUTRIENT SOURCE FOR GULLS 


203 


sites was obtained from M. D. Hocking (personal com- 
munication, Table 2). Fecundity and egg wet mass for 
Pink and Chum salmon (northern mainland coast) 
were obtained from Beacham and Murray (1993). Pink 
Salmon fecundity was 1633 eggs/female and egg wet 
mass was 0.175g; Chum fecundity was 3173 eggs per 
female and egg wet mass was 0.278g. A 1:1 sex ratio 
was used for both Chum and Pink salmon (Heard 
1991; Salo 1991). Total biomass of salmon and eggs 
in the system are summarized for the two watersheds 
in both years in Table 2. 

We calculated guano production for each species 
per day based on Burger et al. (1978) for Kelp Gulls: 
G = 36.lg d'! kg', where G = output (dried) per kg 
body mass per 24 hrs. We adjusted this value to the 
average mass of each gull species. This estimate is con- 
gruent with that of Portnoy (1989) who found that 
Herring Gulls (mass = 1.1kg) produced 39.4 g day’. 


Results 

In both watersheds, six gull species were observed 
to feed on salmon carcasses and eggs: Glaucous-winged 
(Larus glaucescens), Herring (L. argentatus), Thayer’s 
(L. thayeri), California (L. californicus), Mew (L. can- 
us), and Bonaparte’s (L. philadelphia) gulls. The large- 
bodied gulls (Glaucous-winged, Herring, Thayer’s, and 
California gulls) mainly scavenged for salmon car- 
casses and occasionally consumed drifting eggs. Bona- 
parte’s Gulls consumed eggs exclusively and most often 
hovered, “surface-plunging” for eggs. Bonaparte’s 
Gulls also floated and “surface-seized” eggs from below 
the surface. Mew Gulls rarely surface-plunged; most 
of the time they were observed to either surface-seize 
or dislodge eggs from gravels with their feet. From 
behavioural observations of sub-sets of Mew Gulls, 


TABLE |. Average body mass (Dunning 1993), calculated field metabolic rate, and daily consumption of eggs or carcasses 
for large gulls (Glaucous-winged, Herring, Thayer’s, California gulls) Mew Gulls, and Bonaparte’s Gulls. 


Mean Field metabolic Consumption of eggs Consumption of carcasses 
Species body mass (g) rate (KJ/day) (g/bird/day) (g/bird/day ) 
Large gulls 1073.0 1258.9 220.9 569.0 
Mew Gull 403.5 656.2 Lisa 296.6 
Bonaparte’s Gull 212.0 426.6 74.9 192.8 


TABLE 2. Wet mass of salmon carcasses (M. D. Hocking, personal communication) and wet mass of eggs (Beacham and 
Murray 1993) deposited by Pink and Chum salmon at the Clatse and Neekas Rivers. Salmon escapement (Department of 
Fisheries and Oceans) was used to calculate total mass of carcasses and eggs deposited in each watershed in 2002 and 2003. 


Mean carcass Mass Total mass Total mass Total mass Total mass 
wet mass _ of eggs 2002 of salmon of eggs 2003 of salmon of eggs 
Watershed Species (Kg) (g/female) escapement (Kg) (Kg) | escapement (Kg) (Kg) 
Clatse Pink 1.1+/-0.1 285.8 25000 27500 3573 25000 27500 3573 
Chum = 4..24/-0.2 882.1 4300 18060 1897 6000 25200 2646 
Total 29300 45560 5470 31000 52700 6219 
Neekas Pink 1.3+/-0.1 285.8 60000 78000 8574 15000 19500 2144 
Chum = 3..4+/-0.2 882.1 19000 64600 8380 35000 119000 15437 
Total 79000 142600 16954 50000 138500 17580 


204 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE 3. Total consumption of salmon carcasses and eggs during the 60-day study period and proportion of the total salmon 


and egg biomass in the system consumed by gulls. 


Total carcass Total egg 
consumption (Kg) Proportion of total consumption (Kg) Proportion of total 
Watershed Year  original* interpolated** salmon biomass original* interpolated** egg biomass 
Clatse 2002 6318 7931 0.13 - 0.17 1594 1987 0.29 - 0.36 
2003 14221 10419 0.19 - 0.26 2150 1868 0.30 - 0.34 
Neekas 2002 15785 19349 0.11 - 0.14 i fat5 0.07 - 0.09 
2003 26386 24650 0.18 - 0.19 S02 3168 0.17 - 0.18 


*derived from mean of original counts 
**derived from mean of interpolated counts 


approximately 93% consumed eggs and 7% consumed 
carcasses (7 = \11,..s,= 0.19). Gulls obtamed eggs 
buried in the stream gravels, as well as those already 
floating downstream. They were also observed taking 
eggs from carcasses. Feeding intensity of large gulls 
was highest at low tide, when most carcasses in the 
estuary were exposed. 

Gull abundance fluctuated over time at the two 
watersheds (Figure 1). At Clatse River, total daily 
counts of gulls reached a maximum of 1979 (13 Oc- 
tober 2003), of which approximately 45% were large 
gulls. At Neekas River, maximum counts were 3594 
(21 October 2003) of which 64% were large gulls. At 
both watersheds, Glaucous-winged and Herring gulls 
were the dominant species among the large gulls. 
Large gulls increased in abundance over the spawning 
period in both years at both watersheds whereas Mew 
and Bonaparte’s gull abundance was less predictable 
(Figure 1). At the Clatse River, Mew and Bonaparte’s 
gulls peaked in numbers and began to decline in early 
October 2002 and mid-October 2003 (Figure la, b) 
whereas at the Neekas River they did not follow a dis- 
cernable pattern (Figure 1c,d). Total numbers of Mew 
and Bonaparte’s gulls were similar between watersheds, 
whereas greater numbers of large gulls occurred at 
Neekas River. 

Total consumption estimates of salmon carcasses 
and eggs varied between years and watersheds (Table 
3). We estimate that 11% to 26% of total salmon car- 
cass biomass and 7% to 36% of salmon egg biomass 
was consumed by gulls during the study period (Table 
3). Carcass consumption was higher at the Neekas 
than the Clatse River, but proportions of total salmon 
biomass consumed were slightly less at the Neekas 
than the Clatse River. Although egg consumption was 
similar at the two watersheds, substantially higher pro- 
portions of total egg biomass were consumed at Clatse 
compared to Neekas River (Table 3). In general, con- 
sumption was higher in 2003 than 2002 for both water- 
sheds. 

Based on gull counts and body mass, we estimated 
total guano production. At Clatse River, guano output 
ranged from 596 kg to 748 kg in 2002 and 907 kg to 
1192 kg in 2003. At Neekas River, this ranged from 
1201 kg to 1463 kg in 2002 and 2006 kg to 2104 kg in 
2003. Based on observed foraging and resting loca- 


tions of gulls, guano was distributed into multiple 
habitats including the river, riparian zone, estuary, 
and ocean. 


Discussion 

Gulls were significant consumers of the salmon 
resource and used a variety of foraging techniques and 
tissue types. There is an energy trade-off between the 
calorie-rich eggs, which require active searching, and 
the highly available yet low energy-density carcasses. 
Bonaparte’s Gulls, the smallest of the gulls, are well 
adapted to aerial foraging and surface-seizing and com- 
monly feed on insects and zooplankton (Baltz and 
Morejohn 1977; Vermeer et al. 1987; Taylor 1993). 
Their ability to hover above water for extended peri- 
ods of time may facilitate their ability to effectively 
spot and capture eggs in the river. The larger gulls, in 
contrast, with a greater body mass and wing-loading, 
may incur additional energy costs of continuous-flap- 
ping flight which may outweigh the benefits of ob- 
taining the more energy-rich food. These gulls were 
observed on occasion to surface-plunge for eggs, in- 
dicating that at certain times, benefits of capturing eggs 
outweighed energy costs. Although other food sources 
such as benthic invertebrates were available in the 
estuaries, we only observed gulls foraging on salmon 
tissues and eggs. Gulls tend to maximize their utili- 
zation of temporary resources, focusing on localized 
concentrations of prey (Shealer 2002), and it is prob- 
able that when eggs and carcasses are easily available 
on salmon streams, gulls feed solely on this resource. 

Abundance of gulls at the Clatse and Neekas Rivers 
fluctuated over the study period and appeared to cor- 
respond with food availability. Large gull abundance 
increased over time on each watershed in both years, 
corresponding with the accumulation of spawned-out 
salmon on the stream banks and in the estuary. Abun- 
dance of Bonaparte’s Gulls and Mew Gulls, however, 
was not correlated with carcass accumulation. We 
suspect that the rate of egg loss, which is associated 
with spawning density and flooding events, may be 
an important predictor of Mew and Bonaparte’s Gull 
abundance. The two watersheds had similar numbers 
of Mew and Bonaparte’s Gulls despite the higher bio- 
mass of salmon at the Neekas River, indicating that 
comparable quantities of eggs were being lost from 


2005 


Gull Count 


Gull Count 


Day 


CHRISTIE AND REIMCHEN: PACIFIC SALMON AS NUTRIENT SOURCE FOR GULLS 


205 


Gull Count 


Day 


Gull Count 


Day 


FIGURE la-d. Change in abundance of gulls over time (day 1 = 25 August; day 60 = 23 October) for Clatse River: (a) 2002 and (b) 
2003 and Neekas River (c) 2002 and (d) 2003. Dashed lines represent large gulls (Glaucous-winged Gulls, Herring 
Gulls, Thayer’s Gulls, California Gulls), dotted lines represent Mew Gulls, and solid lines represent Bonaparte’s Gulls. 


the two watersheds even though the Clatse River had 
fewer spawning salmon. High stream velocity can 
result in egg loss, causing eggs to be washed out of 
redds after being deposited (Vronskii and Leman 1991). 
This effect may be accentuated at Clatse River where 
recent logging in the headwaters will have lead to 
higher stream velocities and reduced gravel stability. 
Egg loss from salmon redds can also be linked with 
high salmon spawning density, which results in redd 
superimposition and subsequent egg dislodgement 
(Fukushima et al. 1997). It is possible that fluctuations 
in gull abundance were related to timing of migration 
rather than prey abundance. Mace (1983) observed 
aggregations of Bonaparte’s Gulls feeding on juve- 
nile salmonids in the spring and found abundance of 
gulls to be directly related to migration. 

Gulls were major consumers of both salmon car- 
casses and eggs. Our estimates for consumption of car- 
casses at the Neekas and Clatse rivers are conservative 
because our 60-day study period ended before gulls 
had departed from the stream. Extrapolating abundance 
throughout the duration of the spawning period might 
increase consumption by as much as 30%. Our results 
are comparable to those of other studies of gulls feed- 
ing on fish or eggs (Gabrielsen et al. 1987; Haegele 


1993; Bishop and Green 2001). High numbers of egg- 
eating gulls at the Clatse River led to a substantial pro- 
portion (29-36%) of eggs deposited in the system being 
consumed. Only a small proportion of eggs would have 
been dislodged from buried redds; most eggs were 
already floating downstream before capture by gulls. 
It is not unusual for large quantities of eggs to be lost 
from salmon redds; for example, average egg loss rates 
of 48.6% and 56% have been reported for Pink Salmon 
(Eniutina 1972; Heard 1991). In general, higher pro- 
portions of salmon and eggs were consumed in 2003 
than 2002, largely because there was less total salmon 
available in 2003. Between-year differences at the 
Neekas River must be interpreted cautiously, however, 
because of the short study period in 2002. 

Gulls contributed to the cycling of nutrients from 
salmon into terrestrial and aquatic ecosystems through 
guano and feather deposition. Seabird guano has been 
reported to enrich plants in nitrogen and phosphorus 
(Anderson and Polis 1999; Garcia et al. 2002), in- 
crease abundance of terrestrial arthropods (Sanchez- 
Pinero and Polis 2000), and increase primary produc- 
tivity in the intertidal zone (Bosman and Hockey 1986). 
Guano from gulls and other avian scavengers on sal- 
mon streams likely contributes to the nitrogen and 


206 


phosphorous content of otherwise nutrient-deprived 
coastal forests and streams (Waring and Franklin 1979; 
Kiffney and Richardson 2001). In addition, gulls under- 
go an annual molt after breeding (Taylor 1993; Van- 
denbulcke 1989), and their feathers, containing high 
concentrations of mineral elements and energy (Wil- 
liams and Berruti 1978) are shed into the riparian zone, 
stream, and estuary. 

Salmon streams may provide an important food re- 
source for gulls, particularly the smaller species such 
as Mew and Bonaparte’s gulls. Salmon streams offer 
a highly predictable, nutritional and accessible food 
source to gulls dispersing from breeding grounds in 
search of abundant food resources at a time of high 
energy expenditure (feather molt, migration) and high 
juvenile mortality (Burger 1993; Hamer et al. 2002). 
There are roughly 2500 spawning streams in British 
Columbia, many of which attract aggregations of gulls 
during the autumn and winter. Assemblages of gulls 
have been reported to utilize salmon streams in Wash- 
ington (Skagen et al. 1991), Vancouver Island (person- 
al observation), the Queen Charlotte Islands (Reimchen 
1992*) and Alaska (Mossman 1958). The large num- 
bers of gulls observed on the Clatse and Neekas 
Rivers, which have relatively intact salmon runs, are 
representative of an ecological phenomenon that has 
been greatly diluted throughout the Pacific Northwest. 
Gresh and Lichatowich (2000) estimated a 93-95% 
reduction in salmon biomass on the west coast of North 
America over the last century which results reduced 
availability of this food source for gulls and for numer- 
ous other vertebrates species that utilize salmon nutri- 
ents (Cederholm et al. 2000*). The importance of gull 
assemblages to the ecology of coastal terrestrial eco- 
systems is unknown, but gulls have the potential to be 
important nutrient vectors and thus may contribute to 
the primary productivity of nutrient-deprived terrestrial 
systems. 


Acknowledgments 

We thank Alan Burger for his helpful comments on 
the manuscript, Karen Petkau, Morgan Hocking, Janine 
Arnold, Bob Wilkerson, Sara Steinke and Jocelyn 
Akins for assistance in the field, Chris Darimont, 
Raincoast Conservation Society and Larry Jorgenson 
for providing accomodation in the field, Terry Palfrey 
at the Department of Fisheries and Oceans for fish 
escapement data, The Natural Sciences and Engineer- 
ing Research Council of Canada (NSERC), The Friends 
of Ecological Reserves, The David Suzuki Foundation, 
Mountain Equipment Co-op Environment Fund, Bird 
Studies Canada, and Science Horizons Youth Intern- 
ship Program for financial support, and The Heiltsuk 
Nation for allowing this study to take place in their 
territory. 


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Received 12 March 2004 
Accepted 8 March 2005 


Microhabitat Characteristics of Lapland Longspur, Calcarius 
lapponicus, Nests at Cape Churchill, Manitoba 


CLINT W. BoAL!? and Davip E. ANDERSEN? 


'U.S. Geological Survey Texas Cooperative Fish and Wildlife Research Unit, Texas Tech University, Lubbock, Texas, 79409- 
2120 USA 

*Corresponding author: email: clint.boal @ttu.edu 

3U.S. Geological Survey Minnesota Cooperative Fish and Wildlife Research Unit, University of Minnesota, St. Paul, Minnesota 
55108, USA 


Boal, Clint W., and David E. Andersen. 2005. Microhabitat characteristics of Lapland Longspur, Calcarius lapponicus, 
nests at Cape Churchill, Manitoba. Canadian Field-Naturalist 1 19(2): 208-213. 


We examined microsite characteristics at 21 Lapland Longspur (Calcarius lapponicus) nests and land cover types in which 
they occurred in Wapusk National Park, Cape Churchill, Manitoba. Nests were located in four of six physiographic-vegetation 
land-cover types. Regardless of land-cover type, all but one nest was built on a pressure ridge or mound. Nests were built 
midway between the bottom and top of ridges or mounds with steeper slopes than was randomly available. Longspur nests had 
a distinctive southwest orientation (P < 0.001). Longspurs selected nest sites that consisted of comparatively greater amounts 
of shrub species and lesser amounts of moss than were randomly available. Nests were generally well concealed by vegetation 
(mean = 67.0%) and concealment was negatively associated with amount of graminoid species at the nest (P = 0.0005). Our 
nesting habitat data may facilitate a better understanding of breeding Lapland Longspur habitat requirements, and potential 
impacts of habitat degradation by increasing Snow Goose (Chen caerulescens) populations in the study area. 


Key Words: Calcarius lapponicus, Lapland Longspur, Chen caerulescens, Lesser Snow Goose, microhabitat, Cape Churchill, 


nesting habitat, Manitoba. 


The Lapland Longspur (Calcarius lapponicus) 1s one 
of the most common breeding passerines of the Arctic 
(Rising 1996; Gilg et al. 2000; Henry and Mico 2002; 
Hussell and Montgomerie 2002), with a wide distribu- 
tion across well-vegetated tundra landscapes of North 
America and Eurasia (Hussell and Montgomerie 2002). 
However, because breeding by Lapland Longspurs oc- 
curs outside of the Breeding Bird Survey (Dunn 1997), 
little information is available concerning population 
trends (Hussell and Montgomerie 2002). The species 
has experienced substantial population declines in the 
region of Churchill, Manitoba, since the 1930s (Tav- 
erner and Sutton 1934; Jehl and Smith 1970; J. R. 
Jehl, in Hussell and Montgomerie 2002). It is now 
reported to be absent from the area except in small, 
scattered groups along the coast to Cape Churchill 
(Hussell and Montgomerie 2002). 

Understanding a species’ habitat requirements is a 
key element of understanding population trends. Un- 
fortunately, most nesting habitat data for Lapland 
Longspurs are qualitative. Breeding Lapland Longspurs 
are reported as most common in wet tundra, thickly 
vegetated uplands, sedge margins along streams and 
ponds, and hummocks and sedges in marshes (Rod- 
rigues 1994; Rising 1996; Henry and Mico 2002). Nests 
are typically placed in a depression in the ground on the 
side of a bank or hummock (Hussell and Montgomerie 
2002), and consist of a cup of coarse sedges, grasses, 
moss, and roots lined with finer materials such as feath- 
ers, hair, and finer grasses (Watson 1957; Rising 1996; 
Baicich and Harrison 1997). However, few quantitative 


data are available on nest dimensions (Brandt 1943; 
Grinnell 1944; Madsen 1981), orientation (Rodrigues 
1994), vegetation characteristics (Rodrigues 1994), or 
nest concealment of Lapland Longspur nests, all of 
which may relate to nest thermal conditions and con- 
cealment and, thereby, nesting success. Such informa- 
tion is necessary to develop a more complete under- 
standing of the species’ breeding habitat requirements. 
Nesting habitat information may also facilitate a better 
understanding of possible factors leading to local pop- 
ulation trends. Existing data for Lapland Longspurs are 
primarily from the northern portions of the species’ 
breeding range (see Hussell and Montgomerie 2002 for 
review). Fewer published data are available from the 
more southerly extent of the species’ range, and those 
available are dated (Taverner and Sutton 1934; Grinnell 
1944; Jehl and Smith 1970). Here we present data on 
microhabitat characteristics of Lapland Longspur nests 
at Cape Churchill, Manitoba, in June 2003. This coin- 
cided with the peak of nesting activity by the species 
in the area. 


Study Area 

The study area center is approximately 2 km west 
of Hudson Bay in Wapusk National Park, and situated 
35 km east of Churchill, Manitoba and 15 km south- 
east from La Pérouse Bay (58°39'N, 93°11'W). The 
study area is characterized as a tundra biome, consist- 
ing of a series of coastal salt marshes, willow (Salix 
spp.) and spruce (Picea spp.) patches, beach ridges, and 
an inland matrix of large, permanent freshwater lakes, 


208 


2005 


ephemeral freshwater pools, and freshwater sedge 
meadows (Didiuk and Rusch 1979). We characterized 
the landscape into physiographic-vegetation commu- 
nities as described by Henry and Mico (2002). Wet 
Sedge Meadows (WSM) were typically level, hydric 
lowlands with nearly continuous sedge cover and often 
shallow surface water. Graminoid Tundra (GRT) was 
mesic to hydric areas with near-continuous graminoid 
cover and occasional dwarf shrubs (primarily Salix 
spp.). Graminoid/Dwarf Shrub Tundra (GST) consist- 
ed of 75-100% cover of graminoids and moss, with 
herbs and dwarf shrubs on the occasional drier earthen 
hummocks. Common features of the inland matrix are 
low pressure ridges and mounds (i.e., earthen hum- 
mocks) created when surface ice expands during the 
winter and pushes peat upward (Johnson 1987). These 
mounds and ridges support herbs, moss, dwarf shrubs, 
and in lower areas, some graminoids. Some areas con- 
sisted exclusively of these mounds and ridges, and were 
classified as Hummocky Tundra (HT). Dwarf Shrub 
Tundra (DST) occurred primarily on moist but well- 
drained slopes, and consisted of about 50-75% cover 
dominated by dwarf shrubs. Dwarf Shrub/Lichen Bar- 
rens (DLB) were windblown sites at higher elevations 
(e.g., beach ridges), and typically had low (25-50%) 
vegetation cover. All of these descriptions are from 
Henry and Mico (2002), modified to apply to our study 
area. 


Methods 

Intensive nest searches were conducted in one 6-ha 
plot (North Plot) and one 3.7-ha plot (South Plot). The 
North Plot was primarily a HT, but the west border 
also consisted of GRT and WSM. The South Plot was 
primarily GRT with patches of HT and WSM. Inten- 
sive nest searches consisted of walking transects across 
the plot, watching and following longspurs observed 
carrying nest material, prey, or engaged in other behav- 
iors indicative of a nearby nest, or flushing females from 
nests. Transects were closely spaced (e.g., approximately 
20 m apart) but inter-transect distances and linearity 
were variable due to pools and ponds in the search areas. 

The majority of Lapland Longspur nests was not 
located systematically. Rather, many nests were found 
when researchers engaged in other studies inadvertent- 
ly flushed longspurs from nests, or followed longspurs 
observed carrying nesting material or prey. 

A handheld GPS unit was used to record the UTM 
coordinates of each nest site and a flag was placed 10 m 
north of each nest to facilitate relocation. The vegeta- 
tion community at the nest site and the structure (i.e., 
| hummock, pressure ridge) in which the nest was built 
was recorded. The height of the nest above the base of 
| the structure and height of the structure, slope of the 
structure, and orientation of the nest were recorded. 
Nest width and depth were recorded if such measure- 
| ments would not physically disturb the nest or contents. 
| To compare nest orientation to wind patterns, we cal- 


§ 


BOAL AND ANDERSEN: CHARACTERISTICS OF LAPLAND LONGSPUR NESTS 


209 


culated the percent of time the wind was out of the 
north, northeast, east, southeast, south, southwest, west, 
and northwest from daily records at the Nester One 
field station during the period of 3—24 June 2003. 

To assess microhabitat vegetation at nest sites, a 
20-cm by 20-cm frame was centered on the nest and a 
digital photograph was taken from 1.5 m above the nest. 
Each digital image was imported into Adobe Photoshop 
7.0 and overlaid with a l-cm x l-cm grid (100 inter- 
secting points). The vegetation species at each intersec- 
tion was identified using collected samples and pub- 
lished references (Johnson 1987). Proportions of each 
vegetation species and litter were calculated. Identical 
measurements were made at paired random sites situated 
10 m away and in a random direction from each nest. 

We used Adobe Photoshop 7.0 to select the specific 
circular or elliptical area of each nest from the digital 
image. We then resized the image so that the narrowest 
span across the nest was 10 cm. Width of each image 
varied slightly because not all nests were perfectly cir- 
cular. The image was then overlaid with a 0.5-cm by 
0.5-cm grid, resulting in approximately 280-300 inter- 
secting points. The substrate under each intersecting 
point was identified as vegetative cover or nest/nest 
contents. Percent nest cover (1.e., concealment) was then 
calculated as the proportion of all points identified as 
cover vegetation. 

Descriptive statistics for nest dimensions, height, 
slope, orientation, and concealment are provided. We 
used methods described by Zar (1999) to calculate the 
mean bearing and 95% Confidence Interval for nest 
orientation. We used Rayleigh’s test for uniformity to 
test the hypothesis that nest orientation was not ran- 
dom (Zar 1999). Correlation analysis was conducted 
to examine relationships between principal vegetation 
cover types and nest concealment. Rather than signifi- 
cance testing, we compared proportions of vegetation 
species at nest and random sites with 95% Confidence 
Intervals (Johnson 1999). Descriptive statistics reported 
include means and standard deviations. All statistical 
analyses were conducted using program STATISTICA 
6.0. 


Results 

We located 21 active Lapland Longspur nests in the 
study area. Eight of the nests were located in the inten- 
sively surveyed plots, and nesting density in the two 
survey plots was similar (0.83 nests/ha and 0.81 nests/ 
ha). Nearest nesting distance of any two nests was 
39.5 m. Nests were located in 4 of the 6 physiographic- 
vegetation types. Eight nests were in GST (38%), six 
each were located in GRT (28.6%) and HT (28.6%), 
and one was located in DLB (4.8%). Despite the phy- 
siographic-vegetation types where nests were found, 
all but one nest were built on the side or top of a pres- 
sure ridge (38.1%) or mound (57.1%); the exception 
was one nest located in a grass hummock on a beach 
ridge (i.e., DLB). 


210 


60 


f Wind Direction EI) Nest Orientation 


THE CANADIAN FIELD-NATURALIST 


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40 


30 


Percent 


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Vol. 119 


10 


NW 


FIGURE |. Percent of time the prevailing wind blew from the indicated directions and percent of Lapland Longspur nests 
with an orientation in the indicated direction at Wapusk National Park, Cape Churchill, Manitoba, 2003. 


The inner dimensions of Lapland Longspur nests 
were 64.4 mm + 3.9 mm (n = 11) wide and 34.7 mm 
+ 4.6 mm (n = 6) deep. Nests averaged 5.7 cm + 2.2 cm 
above the base of their structures, which was approx- 
imately midway (mean = 55.1% + 17.4%) up the nest 
structure (e.g., pressure ridge). The slope at nests 
(30.1% + 17.8%) appeared to be greater than random 
(17.0% + 13.8%) (difference of means = 13.1% + 25.6%; 
95% Cl = 1.4 % to 24.7 %). Orientation of longspur 
nests was not random (z = 13.252, P < 0.001; Ray- 
leigh’s test for uniformity); longspur nests had a dis- 
tinctive southwest orientation (mean = 208° + 39°; 95% 
CI of 193° to 223°). Forty-eight percent of nests had 
a southwest orientation, and 33% had a southerly ori- 
entation (Figure |). Winds were primarily from the 
north (28%), then west (19%), south (16%) and east 
(13%) (Figure 1). 

Vegetation at the microhabitat scale of the nest 
(400 cm?) consisted primarily of willow (primarily 
Salix arctophila; 21.8%), graminoids (primarily Carex 
aquatilis; 18.9%), moss (10.3%), litter (12.8%), Lap- 
land Rosebay (Rhododendron lapponicum, 10.6%), 
Dwarf Birch (Betula glandulosa; 7.6%), and Bog 
Rosemary (Andromeda polifolia L.; 7.3%). Species 
occurring at lower frequency included Bearberry 
(Arctostaphylos spp.; 2.1%) Black Crowberry (Em- 
petrum nigrum; 2.0%), Dry-ground Cranberry (Vac- 
cinium vitis-idaea; 1.4%), Dwarf Labrador Tea (Ledum 
decumbens; 2.4%), White Mountain-avens (Dryas inte- 


grifolia; 0.3%), and Cloudberry (Rubus chamaemorus; 
0.2%). Based on 95% C.L.s, it appears Lapland Long- 
spurs selected nest sites that had a comparatively greater 
amount of willow, Dwarf Birch, and Lapland Rosebay, 
and lesser amounts of moss (Table 1). 

Lapland Longspur nests were generally well con- 
cealed by vegetation (mean = 67.0% + 22.4%; 95% 
CI = 56.7 — 77.2). Nest concealment appeared to be 
negatively associated with amount of sedge at the nest. 
When an obvious outlier was removed, correlation 
analysis indicated a convincing negative relationship 
between amount of sedge at nest sites and nest conceal- 
ment (r = -0.705, P = 0.0005). This relationship was 
still apparent if the outlier was included (r = -0.474, 
PO 0298). 


Discussion 

Lapland Longspurs appeared to be most abundant 
on wet lowlands and hummocky tundra cover types 
(Williamson and Emison 1971; Hussell and Holroyd 
1974; Montgomerie et al. 1983; Henry and Mico 2002; 
Hussell and Montgomerie 2002). Density estimates 
of the species vary both temporally and spatially. For 
example, average density at Barrow, Alaska, was 75 
pairs/km? over a 19-year period, but was as high as 
200 pairs/km? (Custer and Pitelka 1977). Spatially, den- 
sity estimates from North America range from 17 to 
200 pairs/km? (Hussell and Montgomerie 2002). Al- 
though interpretations of the density estimates in this 


2005 


BOAL AND ANDERSEN: CHARACTERISTICS OF LAPLAND LONGSPUR NESTS 


211 


TABLE |. Mean percent cover (+ SD) of primary vegetative cover types at Lapland Longspur nests (n = 21) and paired sites, 
and mean (+ SD) difference and 95% Confidence Interval for mean difference, Wapusk National Park, Cape Churchill, 


Manitoba, 2003. 


Nest 

Species x SD x 

Willow/Birch 29.4 20.7 Ne Be) 
Sedge 18.9 18.5 18.8 
Moss 10.3 13.8 34.7 
Litter 15.0 8.5 ined 
Lapland Rosebay 10.6 1971 Li 
Bog Rosemary 1.3 8.2 5:9 
Miscellaneous 8.5 12.6 4.6 


study must be made cautiously due to little spatial and 
no temporal replication, Lapland Longspur density at 
the study site (e.g., 82 pairs/km*) does not appear to 
be outside reasonable expectations. In comparison, 
however, the current estimated density of longspurs 
at Churchill was 2.7 to 4.2 pairs/km/? (J. Jehl, personal 
communication). 

Henry and Mico (2002) found Lapland Longspurs on 
Banks Island, Northwest Territories, were most com- 
mon in HT and GST and GRT. Although they did not 
examine nest sites, the pattern of distribution of birds 
seemed to be similar to the landscape types in which 
we found longspur nests. More notable is that regard- 
less of cover type, all longspur nests but one were built 
into pressure mounds or ridges, even in the lower-lying 
graminoid-dominated cover types. 

Qualitatively, Lapland Longspur nests were cups 
woven of dead graminoids and thickly lined with 
feathers and, to a lesser extent, mammal hair which is 
consistent with reports from other studies (Hussell and 
Montgomerie 2002). Inner dimensions of Lapland Long- 
spur nests at Cape Churchill were similar to four nests 
measured at Churchill (mean = 67.5 mm; Grinnell 1944) 
and 22 nests in Greenland (mean = 63.5 mm; Madsen 
1981). Inner dimension of 21 nests at Hooper Bay, 
Alaska, ranged from 76 to 89 mm (Brandt 1943), 
which is inexplicably one to two cm greater than that 
reported elsewhere. Cup depth of longspur nests in our 
study (34.7 mm) was noticeably shallower than the 
42.0 mm reported by Madsen (1981), the 45 mm 
reported by Grinnell (1944), or the 64 to 89 mm 
reported by Brandt (1943). The reason for the differ- 
ences in depth may have to do with sample sizes, 
variability of samples, or with latitude of study. This 
study and that of Grinnell (1944) were near the south- 
ern extent of the species’ breeding range and consisted 
of the smallest sample sizes. The potential for harsher 
summer weather conditions at more northerly latitudes 
(Brandt 1943; Madsen 1981) may have led to longspurs 
building nests with deeper, and thus more thermally 
protected, cups. 

Qualitative descriptions suggest Lapland Longspur 
nests are typically oriented southeast to southwest (Wil- 


Pair 


Difference 95% C.I. for 

SD a SD mean difference 
19.3 9.9 20.8 0.4 to 19.4 
Wie) 0.0 19.6 -8.9 to 9.0 
23.4 -24.4 19.8 -33.4 to-15.3 
11.0 -0.1 10.0 -4.6 to 4.5 
322 9.3 16.3 1.9 to 16.7 
7.4 eS 11.4 -3.8 to 6.5 
7.8 3.9 15.3 -3.1 to 10.9 


liamson 1968, Hussell in Hussell and Montgomerie 
2002). Williamson (1968) suggested Lapland Longspurs 
may orient their nests so that the entrances are oppo- 
site from the prevailing winds and receive greater 
insulation. Also, Lapland Rosebay is more profuse on 
south slopes (J. Jehl, personal communication) and 
longspurs in this study appeared to select that shrub 
as nesting cover. Another explanation is that south- 
facing sides of hummocks may be free of snow earlier 
than other orientations (Hussell and Montgomerie 
2002). On the Arctic Coastal Plain of Alaska, Rodrigues 
(1994) found that Lapland Longspur nests placed on 
the side of a ridge, polygon rim, or tussock (61% of 
nests) had a significant south — southwest orientation. 
However, 39% of the nests he located were placed in 
the open with no detectable orientation. The distinctive 
southwest orientation of nests in our study supports 
the earlier descriptions (Williamson 1968; Hussell and 
Montgomerie 2002) and research results (Rodrigues 
1994) indicating Lapland Longspurs do not randomly 
orient their nests but select for a southerly exposure. 

Lapland Longspur nests are subject to depredation 
by numerous species (Hussell 1974; Hussell and Hol- 
royd 1974; Custer and Pitelka 1977; Fox et al. 1987). 
Nest concealment, especially from visual predators 
[e.g., Parasitic Jaeger (Stercorarius parasiticus), Snowy 
Owl (Bubo scandiacus)]|, may be an important aspect 
of nesting success. Nests in our study area were typi- 
cally well-concealed by overhanging vegetation and, 
in many cases, were quite difficult to locate visually. 
Concealment of nests seemed to be enhanced by herb 
and shrub species, whereas nests dominated by gram- 
inoid cover tended to be more exposed. Unfortunately, 
due to logistical constraints, we were unable to assess 
possible relationships between nest concealment and 
nesting failure. 

Nest microhabitat vegetation usually consisted of 
several species, primarily shrub (e.g., willow, birch), 
woody forbs (e.g., Lapland Rosebay), sedges (e.g., 
Carex aquatilis) and a mixture of herbaceous species. 
It appears that longspurs may have preferred sites with 
shrub and woody forb cover for nesting, but graminoid 
cover was not used disproportionate to availability. 


yAG' 


Similarly, Rodrigues (1994) found shrub and forb 
cover was significantly higher at nest sites than random 
sites, but association with sedge cover was less clear. 

Although Lapland Longspurs appear to be an abun- 
dant species (Hussell and Montgomerie 2002), the 
breeding distribution is north of regions covered by 
the Breeding Bird Survey (Dunn 1997). Monitoring 
the species across its breeding range is impractical, 
but trend information may be gleaned through devel- 
opment of several selected long-term monitoring sites. 
Wapusk NP may provide such a site for long-term 
monitoring for two reasons. First, the study site in 
Wapusk NP is located only 35 km east of Churchill, 
Manitoba, an area that has experienced substantial de- 
clines in Lapland Longspurs (Hussell and Montgomerie 
2002). In contrast to declines at Churchill, survey data 
at the study site in Wapusk NP suggest the species is 
stable or has possibly increased from 1984 to 2000 
(Sammler 2001). Second, the study area is experienc- 
ing increasing amounts of habitat alteration through 
overgrazing by Snow Geese (Chen caerulescens). 
Breeding populations of Snow Geese have experi- 
enced dramatic growth in much of the North American 
Arctic (Ankney 1996), leading to increased densities 
at tundra breeding colony sites and a corresponding 
degradation of vegetation by overgrazing (Kerbes et 
al. 1990; Abraham and Jefferies 1997; Jano et al. 1998). 
As Snow Geese have degraded their preferred foraging 
areas (i.e., salt marshes), they have started foraging 
in inland freshwater sedge meadows (Abraham and 
Jefferies 1997) such as those where this study was 
conducted. Snow Goose-associated habitat alteration 
has led to declines of some species in the Cape 
Churchill area (Abraham and Jefferies 1997). More 
specific to our study site, Lapland Longspurs were 
detected at lower densities in degraded freshwater 
sedge meadows compared to meadows that have not 
been impacted by Snow Geese (Sammler 2001). Thus, 
monitoring Lapland Longspurs at the study site in 
Wapusk NP may provide insight into the potential im- 
pact Snow Goose population increases may have on 
breeding density and reproduction of the species and 
other interior nesting tundra species. 


Acknowledgments 

We thank Murray Gillespie for logistical coordina- 
tion, support, and facilitating our research at the Nester 
One field site while providing the opportunity to en- 
gage in this project. Nestor One is supported through 
the Eastern Prairie Population (EPP) Technical Section 
of the Mississippi Flyway Council. Steve Maxson, 
David Best, Kevin Doherty, Robert Nack, Jeff Dittrich, 
Mike Schroer, Tom Smith, Bruce Golden, and Michelle 
McDowell conducted nest searches across the entire 
study area as part of EPP monitoring efforts. We ap- 
preciate the researchers and staff at the Wapusk Nation- 
al Park and Churchill Northern Research Center for 
support, and the USGS-BRD Minnesota and Texas Co- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


operative Research Units for providing travel funds. 
We thank Geoff Holroyd and Joseph Jehl Jr. for their in- 
sightful and constructive comments on the manuscript. 


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Gilg, O., R. Sane, D. V. Solovieva, V. I. Pozdnyakov, B. 
Sabard, D. Tsanos, C. Zockler, E. G. Lappo, E. E. 
Syroechkovski, and G. Eichhorn. 2000. Birds and mam- 
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Grinnell, L. I. 1944. Notes on breeding Lapland Longspurs 
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Henry, J. D., and M. Mico. 2002. Relative abundance, habitat 
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Hussell, D. J. T. 1974. Photographic records of predation at 
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Hussell, D. J. T., and G. L. Holroyd. 1974. Birds of the True- 
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Hussell, D. J. T., and R. Montgomerie. 2002. Lapland Long- 
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lesser snow goose (Chens caerulescens caerulescens) 


BOAL AND ANDERSEN: CHARACTERISTICS OF LAPLAND LONGSPUR NESTS 


2AS 


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Received 23 April 2004 
Accepted 17 March 2005 


Novel Surface Feeding Tactics of Minke Whales, Balaenoptera 
acutorostrata, in the Saguenay-St. Lawrence National Marine Park 


KATIE J. KUKER, JORDAN A. THOMSON, and URSULA TSCHERTER 


Ocean Research and Education Society (ORES), P.O. Box 117, Les Bergeronnes, Québec GOT 1G0 Canada; e-mail: utscherter 


@ores.org 


Kuker, Katie J., Jordan A. Thomson, and Ursula Tscherter. 2005. Novel surface feeding tactics of Minke Whales, Balaenoptera 
acutorostrata, in the Saguenay-St. Lawrence National Marine Park. Canadian Field-Naturalist 119(2): 214-218. 


Surface feeding behaviours of Minke Whales (Balaenoptera acutorostrata) in the mouth and fjord of the Saguenay River, Québec, 
were documented between June and October 2003. Several novel behaviours associated with gathering prey into dense, near- 
surface aggregations prior to a feeding strike were observed. To our knowledge, these behaviours have not been described in 
detail and may be exclusive to this area or to these individuals. A small number of known Minke Whales show strong site 
fidelity to the Saguenay region, providing an ideal opportunity for the study of foraging behaviour at the individual level. 


Key Words: Minke Whale, Balaenoptera acutorostrata, St. Lawrence Estuary, foraging, surface feeding, specialization, behaviour. 


The Minke Whale (Balaenoptera acutorostrata) 1s 
one of the smallest and most widely distributed baleen 
whales, averaging 7 to 10 m in length and weighing up 
to 9200 kg (Reeves et al. 2002). These whales gener- 
ally undergo annual migrations between low latitude 
breeding grounds in the winter and high latitude feeding 
grounds in the summer. Like other balaenopterids, 
Minke Whales feed by swimming into a concentration 
of small fish or euphausiids mouth agape, then expell- 
ing a large quantity of water through their baleen while 
retaining and swallowing the prey. 

A variety of foraging behaviours have been report- 
ed for this species. In the northeast Pacific, Hoelzel 
et al. (1989) reported two distinct foraging strategies: 
bird-association feeding, in which whales take advan- 
tage of aggregations of shoaling fish beneath feeding 
birds, and lunge feeding, in which whales actively gath- 
er prey prior to a strike that breaches the surface. In 
the St. Lawrence Estuary, Lynas and Sylvester (1988) 
distinguished between patch fishing and line fishing, 
two alternate feeding modes dependent on prey distri- 
bution. Gaskin (1982) and Edds and Macfarlane (1987) 
observed dynamic surface behaviours of Minke Whales 
in the Saguenay-St. Lawrence region but did not des- 
cribe them in detail or address their function. Here, 
we describe and discuss the surface feeding behaviour 
of five well-known Minke Whales in the Saguenay 
Fjord, including several apparently novel manoeuvres. 


Study Area and Methods 

The Saguenay Fjord is a major tributary to the St. 
Lawrence Estuary, running 170 km southeast from 
Lac Saint-Jean to the north shore of the St. Lawrence 
River near Tadoussac, Québec. An 80 m deep sill locat- 
ed 18 km from the mouth of the fjord divides the Sag- 
uenay into distinct inner and outer basins, the outer 
reaching a maximum depth of 250 m (Schafer et al. 
1990). Our study area (Figure 1) consisted of the outer 


basin and mouth of the fjord (48°08'N, 69°43'W), 
which lie within the boundaries of the Saguenay-St. 
Lawrence National Marine Park. Two cetacean species, 
Beluga Whales (Delphinapterus leucas) and Minke 
Whales, are regularly observed in the Saguenay. Larger 
baleen whales, including Fin (Balaenoptera physalus), 
Humpback (Megaptera novaeangliae) and Blue (Bal- 
aenoptera musculus), are occasionally seen at the river’s _ | 
mouth but tend to remain in the more open waters of 
the St. Lawrence Estuary. | 

Data were collected on the foraging behaviour of five 
known Minke Whales (individuals that were photo- 
graphed regularly in the area for at least one full season 
prior to the study) using handheld voice recorders 
during 162 outings between June and October 2003. 
Research outings were conducted on a daily basis 
(weather permitting) aboard | to 3 small (5-7 m) rigid- 
hull inflatable vessels and lasted from 3 to 10 hours. 
Photographs were taken using SLR cameras equipped 
with 300 mm fixed focal lenses on 200 ISO slide film 
to allow identification of individuals based on dorsal 
fin markings and other acquired morphological traits. 
We focused on one identified whale at a time, docu- | 
menting all surface behaviours and taking frequent | 
Global Positioning System (GPS) positions. Duration 
of focal follows used in this study ranged from 15 min- 
utes to several hours and depended largely on the con- 
tinuity of feeding and on tracking conditions. 


Results 

A total of 32.8 hours of surface feeding data were 
collected from the five individuals (M1 — M5) (Figure 2) | 
that showed strong site fidelity to the Saguenay mouth | 
and fjord throughout the season. These whales were } 
observed in the study area on 20, 26, 15, 47 and 31 
days, respectively, from mid-June until mid-October. 
Opportunistic sightings data from the two previous 
seasons showed similar patterns of habitat use by these 


214 


2005 


KUKER, THOMSON, AND TSCHERTER: TACTICS OF MINKE WHALES 


ZA5 


New Brunswick 


FiGuRE |. Map showing location of study site at the confluence of the Saguenay and St. Lawrence Rivers, Québec. 


whales. Sightings of these individuals outside the study 
area were infrequent and occurred mostly near the study 
area boundary. 

In 11 of 43 (25.6 %) focal follows, small fish (ap- 
proximately 10-15 cm) were observed either (a) being 
displaced into the air by a feeding strike; (b) on the 
body surface of the whale during a feeding strike; (c) 
jumping at the surface of the water just prior to a feed- 


ing strike; or (d) being picked up by birds at the feeding 
site prior to or just after a strike. Due to extremely poor 
visibility in the Saguenay we were unable to identify 
the species. When conditions allowed us to approach 
the site of a recent feeding strike, the water was searched 
for scales or dead fish but these efforts proved unsuc- 
cessful. Prey were small shoaling fish but we can only 
speculate that they were most likely juvenile Capelin 


FIGURE 2. Identification photographs of the five individual Minke Whales: a) M1 (lacks dorsal fin markings but has distinct 
bent left fluke tip visible during lateral surface manoeuvres); b) M2; c) M3; d) M4; and e) MS. 


216 


(Mallotus villosus), which has been reported as the 
predominant prey species of Minke Whales in the Sag- 
uenay and in neighbouring areas of the St. Lawrence 
(Gaskin 1982; Simard et al. 2002). Sand Lance (Am- 
modytes americanus) has also been identified as prey 
of Minke Whales in the area (ORES unpublished data) 
and therefore cannot be ruled out. 

The five focal whales used variations of the same 
foraging strategy, which involved actively gathering 
prey at the air/water interface using various combina- 
tions of multi-directional surface manoeuvres followed 
by a feeding strike at or near the surface. Pre-strike 
manoevures that were frequently observed included 
“head slaps”, “chin-up blows” and “exhales on the dive”. 
These behaviours were observed exclusively during 
bouts of surface feeding and were generally performed 
while the whale swam in a tight elliptical pattern fol- 
lowed by a strike at or near the centre of the ellipse. 


Head Slap 

Surfacing dorsal side up, the whale would slowly 
bring its head high out of the water at an angle of 
approximately 30 to 45°. As much as one third of its 
body became visible above the surface with ventral 
grooves not expanded and no water seen being expelled 
from the mouth (Figure 3a). After taking a breath, the 
whale would then quickly and powerfully thrust its 
head toward the water, creating a large splash and an 
audible slap as it struck the surface. Head slaps were 
performed on several surfacings oriented in various 
directions prior to a feeding strike, often at the same 
spot. This manoeuvre was also occasionally performed 
oriented laterally with the whale’s right side striking 
the surface. Head slaps were observed on 336 occa- 
sions (22 of which were lateral) from three whales dur- 
ing 27 focal follows. 99.4% of these were performed 
by M4 and M5 (Table 1). 


Chin-up Blow 

Chin-up blows appeared similar to regular surfacings 
but were more vigorous and performed at a steeper 
angle than regular breaths. As the whale began to sur- 
face, it would raise its head high out of the water, 
breathe, and submerge in a continuous motion with- 
out a slap on the surface. This behaviour was observed 
frequently and was performed while oriented dorso- 
ventrally or laterally (Figure 3b). This manoeuvre was 
observed on 495 occasions (219 of which were lateral) 
from all five individuals during 40 focal follows. Chin- 
up blows were the predominant pre-strike surface 
manoeuvre used by M1, M2 and M3 (Table 1). 


Exhale on the Dive 

Following a respiration, the whale would exhale as 
its blowholes submerged resulting in the displacement 
of a large volume of water into the air (Figure 3c). 
This manoeuvre was typically preceded by a regular 
blow or a chin-up blow and occasionally by a head slap. 
Exhales on the dive were observed on 56 occasions 
and were performed by only M1 and M5 during 11 focal 
follows (Table 1). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Discussion 

The foraging behaviour of the five known Minke 
Whales in the Saguenay is markedly different from that 
observed in adjacent areas of the St. Lawrence River. 
For example, in the Laurentian Channel Head (LCH) 
located approximately 8 km east of the Saguenay mouth, 
Capelin are the primary prey of Minke Whales (Simard 
et al. 2002) and the surface manoeuvres described here 
were not observed in the study period. In the LCH, 
strong tidal currents, a stratified water column and bot- 
tom topography combine to create large areas of upwell- 
ing in which prey are forced to the surface (Simard et 
al. 2002). In contrast, the water in the outer Saguenay 
basin and mouth is well-mixed (Schafer et al. 1990), 
the tidal influence is lower and it appears that these 
whales expend a significant amount of energy gathering 
fish near the surface prior to a strike. 

It is likely that the function of the behaviours des- 
cribed here is to increase prey density at the time of a 
strike. Precisely how this is achieved, however, is un- 
clear and warrants further investigation. It is possible 
that head slaps and exhales on the dive have a combi- 
nation of acoustic, mechanical and visual properties in- 
tended to stimulate an aggregation response in the prey. 
Aggregation has been documented as a common res- 
ponse of shoaling fish to “frightening” visual stimuli 
(Radakov 1973; Tegeder and Krause 1995), however, 
it has not been investigated as a response to mechani- 
cal or percussive acoustic cues such as those produced 
by the Minke Whales in this study. 

Alternatively, it is possible that these behaviours 
temporarily confuse or stun the fish, restricting their 
movement while the whale dives to corral them by cir- 
cling beneath the surface. Weinrich et al. (1992) pro- 
posed this function for lobtail feeding in Humpback 
Whales in the Gulf of Maine. In lobtail feeding, whales 
preying on Sand Lance slap the surface of the water 
with their flukes one to several times while diving and 
follow with the underwater release of bubbles leading 
up to a feeding strike. The response of fish to the release 
of bubbles was examined by Sharpe and Dill (1996), 
who demonstrated that bubbles can be effective in man- 
ipulating (restricting the movement) of groups of Pacif- 
ic Herring (Clupea harengus). 


TABLE |. Pre-strike surface manoeuvres used by five known 
Minke Whales in the Saguenay mouth and fjord. 


ous. Suiiace-Wianoenvie ae 


Individual Chin-up Head Exhale 
blow slap on the dive 
M1 53 0 14 
M2 58 0 0 
M3 180 2 0 
M4 19 154 0 
M5 185 180 42 
Total 495 336 56 


2005 


KUKER, THOMSON, AND TSCHERTER: TACTICS OF MINKE WHALES 


217 


FIGURE 3. Photographs of surface manoeuvres performed prior to a feeding strike: (a) head slap; (b) lateral chin-up blow; 
and c) exhale on the dive. Refer to text for description of these behaviours. 


Other cetacean species have also been observed us- 
ing surface slaps while foraging. For example, Bot- 
tlenose Dolphins (Tursiops sp.) have been reported 
using fluke slaps while foraging for single fish (Con- 
nor et al. 2000) and Dusky Dolphins (Lagenorhynchus 
obscurus) have been reported using them while feed- 
ing on shoaling fish (Wiirsig and Wiirsig 1980). 

Individual Minke Whales in the Saguenay appeared 
to specialize in certain types of feeding strikes (for 
example, strikes oriented laterally or ventral side up) 
and surface behaviours prior to a strike. The five focal 
whales were often observed feeding in close proximity 
to other whales, each using their individual surface 
feeding technique, showing no signs of competition or 
displacement. This consistent variation made it pos- 
sible on many occasions to identify individuals in the 
field by their “signature feeding technique” prior to 
obtaining an identification photograph. Lynas and Syl- 
vestre (1988; Minke Whales) and Weinrich et al. (1992; 
Humpback Whales) have also reported consistent indi- 
vidual variation within a single foraging strategy. These 
studies allude to an individually learned component 
of foraging behaviour, which may result in the devel- 
opment of specialized feeding “styles”. 

Recently, increasing emphasis has been placed on 
the importance of individuality and culture in studies 
of behaviour and ecology (see Bolnick et al. 2003 
and Rendell and Whitehead 2001). Continued studies 
of known Minke Whales in the Saguenay region should 
contribute further valuable insight on these topics. Par- 
ticularly, data collected across seasons on the fidelity 
of these whales to the study area and on the frequency 
and spread of novel behaviours are of interest. Ongoing 
observation of these individuals should lead to an in- 
creased understanding of Minke Whale feeding behav- 
iour and habitat use in the Saguenay, which could have 
important management implications for the Saguenay- 
St. Lawrence National Marine Park. 


Acknowledgments 

Funding for this project was provided by the ORES 
Foundation for Marine Environment Research, Swit- 
zerland. We thank the many volunteer staff and course 
participants of ORES for their help in data collection 
and processing throughout the 2003 season. Many 


thanks also to Larry Dill, Arielle Dylan, Siri Hakala 
and Stephanie Pieddesaux who provided feedback on 
drafts of this manuscript. 


Literature Cited 

Bolnick, D. I., R. Svanback, J. A. Fordyce, L. H. Yang, J. 
M. Davis, C. D. Hulsey, and M. L. Forister. 2003. The 
ecology of individuals: incidence and implications of 
individual specialization. American Naturalist 161: 1-28. 

Connor, R. C., M. R. Heithaus, P. Berggren, and J. L. 
Miksis. 2000. “Kerplunking”: Surface fluke-splashes dur- 
ing shallow-water bottom foraging by bottlenose dolphins. 
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Edds, P. L., and J. Macfarlane. 1987. Occurrence and general 
behavior of balaenopterid cetaceans summering in the St. 
Lawrence Estuary, Canada. Canadian Journal of Zoology 
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Gaskin, D. E. 1982. The ecology of whales and dolphins. 
Heinemann Educational Books, London. 459 pages. 

Hoelzel, A. R., E. M. Dorsey, and S. J. Stern. 1989. The 
foraging specializations of individual minke whales. Ani- 
mal Behaviour 38: 786-794. 

Lynas, E. M., and J. P. Sylvestre. 1988. Feeding techniques 
and foraging strategies of Minke Whales (Balaenoptera 
acutorostrata) In the St. Lawrence River Estuary. Aquatic 
Mammals 14: 21-32. 

Radakoy, D. V. 1973. Schooling in the ecology of fish. Israel 
Program for Scientific Translation and John Wiley and 
Sons, New York. 173 pages. 

Reeves, R. R., B. S. Stewart, P. J. Clapham, and J. A. 
Powell. 2002. National Audubon Society Guide to Marine 
Mammals of the World. Alfred A. Knopf, Inc. New York, 
USA. 

Rendell, L., and H. Whitehead. 2001. Culture in whales and 
dolphins. Behavioral and Brain Sciences 24: 309-382. 
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Fjord: A major tributary to the St. Lawrence Estuary. Jn 
Oceanography of a large-scale estuarine system: the St. 
Lawrence. Edited by M. I. El-Sabh and N. Silverberg. 

Coastal Estuarine Studies 39: 296-320. 

Sharpe, F. A., and L. M. Dill. 1996. The behavior of Pacific 
herring schools in response to artificial humpback whale 
bubbles. Canadian Journal of Zoology 75: 725-730. 

Simard, Y., D. Lavoie, and F. Saucier. 2002. Channel head 
dynamics: capelin (Mallotus villosus) aggregation in the 
tidally driven upwelling system of the Saguenay — St. 
Lawrence Marine Park’s whale feeding ground. Canadian 
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Tegeder, R. W., and J. Krause. 1995. Density dependence 
and numerosity in fright simulated aggregation behaviour 


218 THE CANADIAN FIELD-NATURALIST Vol. 119 


of shoaling fish. Philosophical Transactions of the Royal Wiirsig, B., and M. Wiirsig. 1980. Behavior and ecology of 

Society of London B. 350: 381-390. the dusky dolphin Lagenorhynchus obscurus, in the South 
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in humpback whales, Megaptera novaeangliae. Animal _ Received 6 February 2004 

Behaviour 44: 1059-1072. Accepted 9 May 2005 


First Record of the Plains Minnow, Hybognathus placitus, in Canada 


R. M. SYLVESTER!, S. E. FREELING?, and C. R. BERRY, JR. 


' Montana Fish, Wildlife, and Parks, 475 Fish Hatchery Road, Libby, Montana 59923 USA 

* South Dakota State University, Department of Wildlife and Fisheries Sciences, Box 2140B, NPB 138, Brookings, South 
Dakota 57007 USA 

3 South Dakota Cooperative Fish and Wildlife Research Unit, United States Geological Survey, South Dakota State University, 
Box 2140B, NPB 138, Brookings, South Dakota 57007 USA 


Sylvester, R. M., S. E. Freeling, and C. R. Berry, Jr. 2005. First record of the Plains Minnow, Hybognathus placitus, in Canada. 
Canadian Field-Naturalist 119(2): 219-223. 


Seven Plains Minnows, Hybognathus placitus, Family Cyprinidae, were collected on 11 June 2003 from Morgan Creek, in 
Grasslands National Park, Saskatchewan, Canada. This collection is the first record of the species in Canada and extends the 
northern distribution limit of the species. Of 95 Hybognathus spp. collected at the site, only eight specimens were retained 
for positive identification because of the uncertain status of two conspecifics, the Western Silvery Minnow, H. argyritis, and 
the Brassy Minnow, H. hankinsoni, in Saskatchewan. Our findings should stimulate additional sampling to assess the identi- 
fication and status of Hybognathus spp. in southwestern Saskatchewan. Accurate field identification of Hybognathus spp. 
remains an issue and collection of all specimens is recommended to accurately identify members within the genus. 


Key Words: Plains Minnow, Hybognathus placitus, Hybognathus spp., first record, range extension, Morgan Creek, Rock 


Creek, Saskatchewan, Canada. 


The Plains Minnow, Hybognathus placitus, Family 
Cyprinidae, is a small (< 15 cm; Baxter and Stone 
1995), short-lived (Taylor and Miller 1990; Pflieger 
1997), sexually dimorphic (Ostrand et al. 2001), her- 
bivorous species (Page and Burr 1991; Hesse 1994). 
Reproduction coincides with high or receding flows 
in the spring and summer, after which many adults 
die (Sliger 1967; Taylor and Miller 1990; Pflieger 
1997; Platania and Altenbach 1998). The Plains Min- 
now inhabits slower waters and side pools of larger, 
turbid streams, mainly west of the Missouri River from 
Montana and North Dakota, south to central Texas 
(Baxter and Stone 1995; Pflieger 1997). 

Similar morphology and variation in meristics with- 
in Hybognathus spp. has made many aspects regarding 
identification, taxonomy, and phylogeny difficult, but 
efforts have been made to clarify sources of confu- 
sion throughout their range (Niazi and Moore 1962; 
Al-Rawi and Cross 1964; Schmidt 1994; Scheurer et 
al. 2003). Efforts concerning identification have proven 
beneficial, especially when morphologically similar 
species of the genus Hybognathus coexist. The shape 
of the basioccipital bone, dorsal fin profile, and number 
of apical scale radii are the most definitive methods to 
distinguish among members of the genus. The basioc- 
cipital bone appears peg-like in H. placitus compared 
to the broad process of the Western Silvery Minnow, 
H. argyritis, and the Mississippi Silvery Minnow, H. 
nuchalis, and the straight-edged process of the Eastern 
Silvery Minnow, H. regius (Page and Burr 1991). The 
rounded dorsal fin profile and number of apical scale 
radii (about 20) can be used to separate the Brassy Min- 
now, H. hankinsoni, from other Hybognathus spp. that 


have a pointed dorsal fin profile and from 8-14 apical 
scale radii (Baxter and Stone 1995; Pflieger 1997). 
When H. placitus and H. argyritis coexist, they often 
segregate ecologically, with H. placitus inhabiting the 
main channel and H. argyritis inhabiting protected 
backwaters and channel border habitats (Pflieger 
1971; Welker 2000). 

The Plains Minnow is listed in Colorado, Kansas, 
and North Dakota as a species of concern but has no 
status in Alberta, Saskatchewan, Montana, Wyoming, 
South Dakota, Nebraska, Iowa, and Missouri (Loomis 
1997). Plains Minnow have been collected from the 
Rock Creek drainage (R. Lott, Montana Fish, Wildlife, 
and Parks, Glasglow, unpublished data, 2003), the 
Frenchman River drainage (Sylvester 2004), and sev- 
eral tributaries to the Missouri and Yellowstone Rivers 
(Gould and Brown 1966) in Montana. Plains Minnows 
have never been reported in Canada but absence of the 
species in Saskatchewan is likely due to lack of sam- 
pling effort (R. E. Jensen, Saskatchewan Environment 
and Resource Management Swift Current personal 
communication, 2003). Modifications such as dams 
that cause changes in the hydrologic regime, increases 
in water clarity, and species introductions have caused 
the species to be extirpated from many systems (Cross 
and Moss 1987; Pflieger and Grace 1987; Tomelleri and 
Eberle 1990; Hesse 1994). However, the species often 
persists where relatively undisturbed habitat conditions 
exist in the upper Missouri River basin. 

The purpose of this study is to report the first known 
finding of the Plains Minnow, H. placitus, in Canada. 
Information was collected in the Rock Creek drainage, 
Saskatchewan, as part of an accuracy assessment of 


219 


220 


the fish distribution models for the Aquatic Gap Analy- 
sis Program of the United States Geological Survey 
(Wall et al. 2002*). 


Materials and Methods 
Study Area 

The Rock Creek drainage is located in southwestern 
Saskatchewan and north-central Montana (Figure 1). 
Rock Creek flows southerly and is a direct tributary to 
the Milk River, which flows into the Missouri River 
below Fort Peck Reservoir, Montana. Rock Creek flows 
through mixed-grass prairie that contains species such 
as Needle-and-thread Grass (Stipa comata), Blue Gram- 
ma (Bouteloua gracilis), Western Wheat Grass (A gro- 
pyron smithii), sagebrush (Artemisia sp.), Grease- 
wood (Sarcobatus vermiculatus), Prickly Pear Cactus 
(Opuntia polyacantha), Buckbrush, (Symphoricarpos 
accidentalis), willow (Salix sp.), Thorny Buffalo Berry 
(Shepherdia azgenteas), Trembling Aspen (Populus 
tremuloides), and Manitoba Maple (Acer negundo), 
(Parks Canada 2002*). Major anthropogenic features 
such as dams have not altered the drainage in Saskat- 


THE CANADIAN FIELD-NATURALIST 


MILK RIVER 


Vol, 119 


chewan but farming and livestock grazing have impact- 
ed the local landscape and some streams (Parks Cana- 
da 2002*). Three sites were sampled from 10 June 
2003 to 11 June 2003 (Figure 1). 


Site Selection and Fish Sampling 

Site selections in the Rock Creek drainage were 
based on access to streams via public road crossings 
and contacts with personnel from the Grasslands 
National Park office in Val Marie, Saskatchewan. A 
Smith-Root Model LR-24 backpack electrofisher was 
the primary fish sampling gear and no block nets were 
used. Settings were adjusted outside of the reach until 
fish were sufficiently stunned and vulnerable to dip 
netting. Electrofishing proceeded upstream in a zigzag 
pattern and sampled all available habitat types. Fishes 
captured while electrofishing were held in a live cage, 
while additional sampling with a bag seine (5 or 10 m 
x 1.2 m, 4.8 mm mesh) was performed. Fishes collect- 
ed using the seine were placed in a separate live cage. 
All fishes were then identified to species and counted 
by gear type. Specimens that could not be identified 


CANADA 
USA 


we 


ae “te. ee. OX 
5 


\ 
ol . 


if 
, rf 
; pees: “Ne 
| 
Ye 
pom. 
gh 
20 cr 40 50 Kilometers 


FIGURE 1. Location of the Rock Creek watershed in Canada and the United States. The dashed line indicates the internation- 
al boundary between Canada and the United States and the points with corresponding numbers represent the 2003 sam- 


ple locations within the Rock Creek watershed. 


2005 


SYLVESTER, FREELING, AND BERRY: PLAINS MINNOW IN CANADA 


pp | 


TABLE |. Number of fish collected by species at each of three sites sampled in the Rock Creek drainage, Saskatchewan, 


Canada, from 10 to 11 June 2003. 


Species Common Name Site 1 Site 2 Site 3 
Catostomus commersonii White Sucker 36 20 55 
Couesius plumbeus Lake Chub 5 dp 118 
Culea inconstans Brook Stickleback 10 0 67 
Etheostoma exile Iowa Darter I 0 0 
Hybognathus hankinsoni Brassy Minnow 0 l 20 
Hybognathus placitus Plains Minnow 0 fi 0 
Hybognathus spp. Unknown 0 87 127 
Margariscus margarita Pearl Dace 33 69 347 
Phoxinus eos Northern Redbelly Dace 35 0 197 
Pimephales promelas Fathead Minnow 64 ig | 107 
Rhinichthys cataractae Longnose Dace i i 6 


in the field were retained following collection permit 
stipulations, fixed in 10% formalin, and later identified 
in the laboratory at South Dakota State University, 
Brookings, South Dakota, United States. 


Measurements, Meristic Counts, Sex, and Maturity 

Hybognathus placitus were measured to the nearest 
millimeter for total and fork length and weighed to the 
nearest gram. Meristic counts were performed follow- 
ing standard methods (Al-Rawi and Cross 1964). Scales 
above the lateral line were counted across the back 
using the second row of scales in front of the dorsal fin 
insertion. Scales below the lateral line were counted 
around the belly using the second row of scales in 
front of the pelvic fins. Scales were collected below 
the dorsal fin and above the lateral line for approxi- 
mate apical scale radii count (i.e., <15 or closer to 
20). Sex and maturity of H. placitus vouchers were 
determined in the lab by dissection and microscopic 
examination of gonads. The shape and profile of the 
basioccipital process was also examined after dissec- 
tion. Voucher specimens were added to the fish col- 
lection at South Dakota State University. 


Results and Discussion 

A total of 1656 fish representing 10 species were 
collected in our survey of the Rock Creek drainage in 
Saskatchewan. Pearl Dace, Margariscus margarita, 
Fathead Minnow, Pimephales promelas, and Northern 
Redbelly Dace, Phoxinus eos. dominated the catch 
(Table 1). No Hybognathus spp. were collected at site 
1 (49904'15.6"N 106°31'49.9"W). A total of 222 speci- 
mens were tentatively identified as Hybognathus spp. 
in the field from sites 2 (49°00'32.2"N 106°39'51.9"W) 
and site 3 (49°10'27.9"N 106°33'57.7"W). Eight Hybo- 
gnathus specimens were retained for positive identifi- 
cation from site 2 and the remaining specimens (N = 87) 
were released because of the uncertain status of con- 
specifics, H. argyritis and H. hankinsoni. Based on 
communications with provincial fisheries personnel 
(R. E. Jensen, Saskatchewan and Resource Manage- 
ment, personal communication), the field identity of 


voucher Hybognathus spp. specimens from site 2 was 
believed to be H. argyritis. Of the eight Hybognathus 
spp. vouchers collected from site 2, 1 was identified as 
H. hankinsoni and 7 as H. placitus, a previously undoc- 
umented species in Saskatchewan and Canada. The 
pointed dorsal fin profile, peg-like shape of the basioc- 
cipital bone, and the number of apical scale radii (<15) 
were used to identify Hybognathus placitus specimens. 
Hybognathus placitus specimens (post fixation) ranged 
from 46 to 91 mm total length, from 44 to 84 mm fork 
length, weighed from 0.7 to 5.8 g, and included one 
mature, gravid female (Table 2). Meristic counts were 
similar to specimens from Wyoming and South Dakota 
(Al-Rawi and Cross 1964). No specimens were iden- 
tified as H. argyritis. Known H. hankinsoni were col- 
lected at site 3 (males in color) and unknown vouchers 
from site 3 believed to be H. hankinsoni were verified 
as H. hankinsoni in the lab. 

Habitat at site 2 appeared to represent the preferred 
habitat of the H. placitus. The majority of the site was 
run and pool habitat, water velocities were slow (< 0.5 
m/s), substrates were generally small (< 2.0 mm), 
and water was turbid. Site 1 contained predominately 
cobble substrate, which is not the preferred habitat of 
H. placitus (Pflieger 1997). Site 3 had some boulder 
and cobble present due to a culvert and road crossing, 
was spring fed, had high water clarity, and contained 
bog-like vegetation and cattails (Typha spp.). Habitat 
conditions at site 3 were more suitable for the Northern 
Redbelly Dace, which were the second most abundant 
species at the site. Mean wetted width of the sites 
ranged from 2.26 to 3.24 m, total dissolved solids 
ranged from 740 to 1270 (S, temperature from 13.9 to 
16.8°C, salinity from 0.3 to 0.6 ppt, specific conductiv- 
ity from 699 to 1150 (S, pH from 8.4 to 8.9, and dis- 
solved oxygen ranged from 7.6 to 10.7 ppm. Riparian 
vegetation was a mixture of grasses, sedges, and shrubs 
at all three sites. 

This collection of seven Plains Minnow, H. placitus, 
in Morgan Creek, Saskatchewan represents a new spe- 
cies record in Canadian waters and extends the northern 


aad 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE 2. Post fixation measurements, meristic counts, sex, and maturity of H. placitus specimens collected from Morgan 
Creek, Saskatchewan on 11 June 2003. An asterisk identifies characteristics used to identify H. placitus specimens from 


other members of the genus. 


Characteristic 1 2 
Sex Unknown Female 
Maturity Immature Mature 
Total length (mm) 46 76 
Fork length (mm) 4-4 70 
Weight (g) 0.7 4.1 
Lateral line scales 36 38 
Scales above lateral line 13° 13 
Scales below lateral line 19) 18 
Pectoral fin rays 15 16 
Anal fin rays 8 8 
Dorsal fin rays 8 8 
Dorsal fin profile* Pointed Pointed 
Basioccipital process shape* — Peg-like Peg-like 
Apical scale radii* <15 <15 


distribution limit of the species. Hybognathus argyritis, 
H. hankinsoni, H. nuchalis, and H. regius have all been 
documented in Canada (Willock 1969; Scott and Cross- 
man 1973; Bishop 1975; Harbicht et al. 1988; Page and 
Burr 1991; Nelson and Paetz 1992; Houston 1998) and 
Hybognathus argyritis is listed as threatened in Alberta. 
- Although collection of H. placitus from Morgan Creek 
is not surprising based on other collection locations 
within the Rock Creek drainage and the adjacent 
Frenchman River drainage, it is the first known record 
in Canada. Presence of the species is likely due to the 
relatively undisturbed conditions in that portion of the 
Rock Creek watershed. After our survey in 2003, more 
detailed examination of H. argyritis museum speci- 
mens previously collected in Saskatchewan resulted 
in changes of their identity to H. hankinsoni (K. M. 
Murphy, Saskatchewan Environment and Resource 
Management, Swift Current, personal communication). 
Therefore, the presence of H. argyritis in Saskatchewan 
has not been verified to date. 

Collection and preservation of all Hybognathus 
specimens may be the only way to determine the true 
identity and presence of Hybognathus spp. in Canada 
because accurate field identification is difficult. We 
recommend further investigation into the distribution, 
abundance, population characteristics, life history, 
and identification of Hybognathus spp. in Saskatchewan 
and suggest addition of H. placitus to the species list of 
both Saskatchewan and Canada. Although the species 
is common elsewhere, protection under the recently 
passed Species at Risk Act in Canada may be war- 
ranted after further investigation into the population 
status of the species and other members in the genus. 


Acknowledgments 
We would like to acknowledge R. Jensen for assist- 
ing us in the field, with landowner contacts, and for 


Specimen Number 


5 4 5 6 is 
Male Male Male Male Male 
Mature Mature Mature Mature Immature 
87 9] 86 91 71 
79 84 79 84 66 
5.8 5.6 4.4 5d P| 
38 37 39 38 39 
13 13 13 13 13 
i) 15 15 16 18 
16 16 iS 16 15 
8 8 8 8 8 
8 8 8 8 8 
Pointed Pointed Pointed Pointed Pointed 
Peg-like Peg-like Peg-like Peg-like Peg-like 
<b <15 <5 <15 <15 


comments on this manuscript. We also acknowledge 
S. Wall for generating maps in this manuscript and for 
field assistance. We would also like to thank K. Murphy, 
N. Morey, C. Hoagstrom, and J. Duehr for comments 
on this manuscript. Additional thanks go to Grasslands 
National Park personnel for assistance and coordination 
of sampling locations and to R. G. Bramblett for ver- 
ification of Plains Minnow specimens. 


Documents Cited (marked * in text) 

Parks Canada. 2002. Grasslands National Park of Canada 
Management Plan, October 2002. 

Wall, S.S., C. J. Kopplin, B. L. Kopplin, J. A. Jenks, and 
C. R. Berry, Jr. 2002. Expanding South Dakota Aquatic 
Gap Analysis to the Upper Missouri River Basin. Pages 
40-43 in GAP Analysis Bulletin Number 11. United 
States Department of the Interior and United States Geo- 
logical Survey, Moscow, Idaho. 


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Cross, F. B., and R. E. Moss. 1987. Historic changes in fish 
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a 


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Received 25 May 2004 
Accepted 6 June 2005 


Identification and Status of the Introduced Black Pine, Pinus nigra, 
and Mugo Pine, Pinus mugo, in Ontario 


PAUL M. CATLING 


Biodiversity, National Program on Environmental Health, Agriculture and Agri-food Canada, William Saunders Building, 
Central Experimental Farm, Ottawa, Ontario, Canada K1A 0C6; e-mail catlingp @agr.gc.ca 


Catling, Paul M. 2005. Identification and status of the introduced Black Pine, Pinus nigra, and Mugo Pine, Pinus mugo, in 
Ontario. Canadian Field-Naturalist 119(2): 224-232. 


Recent reference texts and other sources are contradictory regarding the spread of Pinus nigra and Pinus mugo from cultivation 
in Ontario. Both species have reproduced from plantings along roadsides but neither has been observed to occupy habitats 
in Ontario to the substantial reduction or exclusion of native species, or to substantially invade natural habitats. The more 
widely planted Black Pine has been recorded spreading at 24 localities throughout the eastern part of southern Ontario. 
Mugo Pine is reported spreading at 18 locations. Although evidently much less aggressive than Scots Pine (Pinus sylvestris), 
both Black and Mugo pines have a potential for negative impact on biodiversity in dry, rocky or sandy habitats, especially in 
connection with extensive plantings. A key for the identification of two-needle pines is included. Both P. nigra and P. mugo 
are highly variable and reported to hybridize extensively with other species. 


Key Words: Black Pine, Austrian Pine, Pinus nigra, Mugo Pine, Pinus mugo, Scots Pine, Pinus sylvestris, spreading, invading, 


alien, naturalized, woody plants, Ontario. 


Introduced woody plants are increasingly a source of 
questions relating to identification, status, distribution, 
and general biology. In many cases, authoritative an- 
swers to the questions are not readily available. Accu- 
rate information is important because invasive alien 
woody plants are a major threat to Canadian native 
biodiversity (Catling 1997). Some are also of impor- 
tance with respect to agriculture, horticulture and/or 
forestry. The introduced Black Pine (Pinus nigra 
Arnold) and Mugo Pine (Pinus mugo Turra) provide a 
good example. They are both important, particularly in 
landscaping, but their current status, potential to spread 
from cultivation and their distinguishing features are 
not well known. 


Status of Black Pine and Mugo Pine in 
North America 

A tree to 30 m high with a rounded crown, Black 
Pine, also called Austrian Pine, is native to Eurasia 
and north Africa from Spain and Morocco east to 
eastern Turkey and north to Austria (Dallimore et al. 
1966; Farjon 1984; Mirov 1967, map). It has been 
planted in North America for dune stabilization, for- 
estry and horticultural purposes (Burns and Honkala 
1990), and is currently one of the most common intro- 
duced ornamentals in the United States (van Haver- 
beke 2002*). 

Kral (1993) does not include Black Pine in the key 
and species accounts in his recent work on North Amer- 
ican pines, but he does make reference to it being 
naturalized in Illinois and notes characteristic features 
under general notes on the genus. Gleason and Cron- 
quist (1991) do not include Black Pine in their key, 
but allude to it as a local escape under Red Pine. Farrar 
(1995) indicates that it is not spreading in Canada. 


Black Pine is actually much more widely escaped than 
these standard references suggest (Figure 1). It is nat- 
uralized throughout the Great Lakes states (e.g., Burns 
and Honkala 1990; Swink and Wilhelm 1994; Mitchell 
and Tucker 1997; Parfitt and Wade 2000; Leege and 
Murphy 2001) and in New England (e.g., Haines and 
Vining 1998, van Haverbeke 2002*), in the Pacific 
Northwest (Petrides and Petrides 1998) and it has been 
listed as an invading species in Canada (Catling 1997). 

Mugo (Mugho) Pine, also known as Mountain Pine, 
is native to the mountains of central Europe and the 
Balkan peninsula. It is not included in the North Amer- 
ican flora by Kral (1993), presumably because it had 
not been reported as spreading. It is widely used in 
landscaping and in stabilization of steep slopes. How- 
ever, it was listed as spreading in Canada (Catling 
1997) based on observations in Ontario. 

The Ontario plant list (Newmaster et al. 1998) gives 
the status of both Black Pine and Mugo Pine as “orna- 
mental’, there defined as “‘plants that have escaped from 
gardens” (page 18), but this category has evidently 
been applied to plants that persist after cultivation in 
Ontario, as well as to those that spread. The authors 
were unaware of any case of these pines spreading 


(personal communication). Thus the 5-20 occurrences | 


suggested by the provincial rank of “SE2” and “SE1” | 


for Black Pine and Mugo Pine (respectively) refer to | 


populations persisting after cultivation since the num- 
ber of locations of plantings in Ontario is certainly in — 
the many hundreds for both species. 

Even when included in texts the two species have 
not been adequately compared with similar species. 
For example, both Japanese Black Pine (P. thunbergii 
Parlatore) and Japanese Red Pine (P. densiflora Sie- 
bold & Zuccarini) have escaped from cultivation in 


224 


2005 


CATLING: BLACK PINE AND MUGO PINE IN ONTARIO 


22 


: 5a NOR, « as 


Ficure 1. A ten-year-old Black Pine (Pinus nigra) spread from a 30-year-old roadside planting along highway 7 in Hastings 


County, Ontario, Canada. Photo by P. M. Catling. 


northeastern North America, and both could be con- 
fused with Black Pine but are not included in available 
keys. 

A continuing assessment of the impact of these two 
alien pines in Ontario is desirable since they are current- 
ly being planted widely in North America. For example 
several million trees of Black Pine are produced 
annually in the United States (van Havenbeke 2002*). 
Assessment of impact is dependent upon a better under- 
standing of both status and identification. The follow- 
ing work addresses these needs. 


Methods 

Literature on Black and Mugo Pines was reviewed 
in order to place occurrence and status within Ontario 
in a global context, and to provide information for 


identification. An identification key was prepared based 
on published studies and examination of specimens. 

Locations in Ontario where young trees of Black 
Pine and Mugo Pine were growing near older plantings 
were recorded. Voucher specimens were collected and 
deposited in the Agriculture and Agri-Food Herbarium 
(DAO) in Ottawa. 

Curators of various herbaria with significant On- 
tario collections including CAN, DAO, HAM, OAC, 
QK, TRT, TRTE, UWO, and WAT (acronyms from 
Holmgren et al. 1990), were contacted with a request 
to examine their holdings and databases for informa- 
tion on Black Pine spreading from cultivation. Field 
botanists and natural resource biologists were also 
contacted to find out if they had observed escaping 
populations. 


226 


Results and Discussion 
(1) Beneficial and detrimental aspects 
Beneficial Aspects 

Both Black and Mugo Pines are widely recognized 
as valuable ornamental plantings due to resistance to 
pollution and high tolerance of de-icing salt spray along 
roadsides. Black Pine may be the most pollution-toler- 
ant species of pine (Earle 2001*). Both species have 
also been recommended for use as windbreaks and as 
bioindicators of environmental pollution (Micieta and 
Murin 1998). Black Pine has also been recommended 
as a useful tree for tracking climatic change on a local 
scale (Levanic 1999; Collins et al. 2000). 


Forestry Importance 

Black Pine is the primary host of Diplodia tip rust 
in parts of the Great Lakes region (Vujanovic et al. 
2000; Michigan State University http://www.msue.msu. 
edu/msue/imp/mod03/01701195.html). This rust infects 
native pines and other conifers. Among the recommend- 
ed control measures is the use of native plantings in- 
stead of exotic plantings. 


Biodiversity Importance 

In Allegan County, Michigan, where 26 000 Black 
Pine trees were planted in a dune system between 1956 
and 1972, and where the trees are now reproducing 
and spreading, the pine stands have been associated 
with a reduced cover of dune vegetation and depressed 
species richness (Leege and Murphy 2001). There was 
also evidence for modification of dune habitats at this 
site and the stands of introduced pines appeared to be 
functionally different from native tree stands. “Repro- 
duction and naturalization of this tree in large num- 
bers” has occurred in Illinois Beach State Park, Lake 
County, Illinois (Swink and Wilhelm 1994). An impov- 
erished native seedbank has been reported in soils under 
Black Pine planted in natural dolomite grasslands in 
Hungary (Csontos et al. 1997). 

Alien conifers replacing natural plant communities 
are a major problem in New Zealand (Hunter and Dou- 
glas 1984) where a recent study found that control of 
Black Pine required greater herbicide applications than 
were required to control other spreading conifers (Lang- 
er 1992). Both Mugo Pine and Black Pine are alien 
species of major concern to the conservation of natural 
habitats in New Zealand and extensive control pro- 
grams are in effect (e.g. New Zealand Department of 
Conservation 2002*). 


(2) General Survey Results 

Both Black Pine and Mugo Pine were found to have 
spread from cultivation at a number of sites (Figures 
2 and 3). There was no recent evidence of planting or 
cultivation in any of these areas. Most of these sites 
were old field or woodland edge habitats along roads. 
At many locations the young trees were of different 
ages from | to 20 years. They were not equidistant, but 
in patches and/or close together and near the putative 
parent. These observations support the conclusion that 
they had spread naturally from the plantings. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Remarkably, there was not a single herbarium speci- 
men of either Black Pine or Mugo Pine in any of the 
collections surveyed that had a label suggesting escape. 
The potential for trees to be invasive is not immediately 
apparent because of the time that it takes to reach repro- 
ductive maturity. In the case of Black Pine, trees can 
reproduce as early as 15-20 years of age, or can delay 
until much later (Vidakovic 1974). It appears that spread 
of Black Pine in Ontario has occurred only over the past 
few decades. Good seed production in Black Pine occurs 
every 3-5 years (Kerr 2000). The delay as well as peri- 
odicity in ample seed production, may partially explain 
the lack of herbarium collections. 

Nevertheless, the lack of observation, considering 
that many established trees that had evidently spread 
from plantings were over 10 years of age, suggests that 
invasive species are easily overlooked, and that the 
manpower available to document invasion is limited. 
Although invasive plants are a serious ecological prob- 
lem, much more effort is currently devoted to docu- 
mentation of rare native species. 


(3) Occurrence of Black Pine in southern Ontario 

Spread of Black Pine, although not frequent, has 
occurred over an extensive area of southern Ontario 
(Figures | and 2). The vouchers and most other trees 
examined were referable to var. nigra, which is to be 
expected since most of the North American plants orig- 
inate from seeds collected in Austria (van Haverbeke 
2002*). Those records for which vouchers are available 
include: 

ONTARIO: HASTINGS: Hwy 7 at Madoc turnoff, 44.5250°N, 
77.4176°W, old trees of similar age (30 years) and 20 young 
trees 2-8 years old, 22 Oct. 2001, P M. Catling 2001-4 (DAO); 
Hwy 7, E of Madoc turnoff, 44.5036°N, 77.5071°W, at this 
site there were about 30 old trees of similar age (30 years) 
and 10 young trees 1-8 years old, 22 Oct. 2001, P M. Cat- 
ling 2001-5, (DAO); Hwy 7 near Black River, 44.5395°N, | 
77.371 1°W, 6 year old tree escaped from roadside plantings, / 
22 Oct. 2001, P M. Catling (DAO). Hwy 7 near Madoc, | 
44.4943°N, 77.6405°W, at least 50 young plants, many seed- 
lings, within 30 m of a large planted tree, 22 Oct. 2001, Pj 
M. Catling 2001-10 (DAO). OTTAWA-CARLETON: Hwy 417 | 
and Boundary Road, | km S of Vars, 45.3379°N, 75.3447°W, 
16 May 2002, P. M. Catling & V. R. Brownell (DAO). UNITED 
COUNTIES OF STORMONT, DUNDAS, AND GLENGARRY: S side | 
of hwy 401 W of Cornwall, 45.0556°N, 74.8006°W, two 3- 
year-old seedlings under seven 30 year old planted trees, 29 
Sept. 2001, P M. Catling (DAO); Hwy 43, 3 km W of } 
Avonmore, 45.1613°N, 75.0214°W, 15 May 2002, P M. | 
Catling & V. R. Brownell (DAO). 

In addition to these vouchered observations, 17 sight | 
records of young trees spread from cultivation are plot- | 
ted on the accompanying map (Figure 2). | 


(4) Occurrence of Mugo Pine in southern Ontario 

Although it is widely cultivated and capable of re- | 
producing when only 10 years of age, there are relative- | 
ly few records of Mugo Pine escaping from cultivation | 
in Ontario (Figure 3). All three subspecies are repre- 
sented by escapes in Ontario. Those records for which | 
vouchers are available include: | 


2005 


CATLING: BLACK PINE AND MUGO PINE IN ONTARIO pape 
76° TA 
i 
100 Kilometers 
z 
+ 
LS 
sar 
| 8 
Pinus nigra (escaped) ‘ 
@ herbarium collections 
A sight records 


84° 82° 80° 


78° 76° 


FiGuRE 2. Central and eastern portions of southwestern Ontario showing locations where Black Pine (Pinus nigra) has 
escaped from cultivation. Escaped occurrences supported by herbarium specimens at DAO, Agriculture and Agri- 
food Canada, Ottawa, are indicated with a solid dot. Sight records of the author are indicated by solid triangles. 


OnrTARIO: HasTINGs: Hwy 7 east of Marmora, 44.4943°N, 
77.6566°W, escaped from plantings, 22 Oct. 2001, P.M. Cat- 
ling 2001-20 (DAO sub subsp. mugo); Hwy 7 east of Marmo- 
ra, 44.4943°N, 77.6566°W, from a 10 year old escaped plant 
with cones with apophysis hooked and recurved, accompa- 
_ nied by seedlings beneath a plant referable to subsp. unci- 
— nata, 22 Oct. 2001, P. M. Catling 2001-11 (DAO sub subsp. 
__uncinata); 13 km east of Marmora along Hwy 7, 44.5033°N, 
_ 77.5116°W, this branch collected from a small sapling 4 m 
tall, evidently escaped from planted shrubs 5 m tall within 
_ 20m, many young plants 1-10 years old, 22 Oct. 2001, P. M. 
Catling 2001-8-2 (DAO sub nothosubsp. rotundata); Hwy 7 
_ east of Marmora, 44.4943°N, 77.6566°W, % year old plant, 
_ escaped from roadside plantings, 22 Oct. 2001, P M. Catling 
— 2001-11, (DAO sub nothosubsp. rotundata); UNITED COUNTIES 
_ OF LEEDS AND GRENVILLE: 2 km NE of Brockville, 44.6263°N, 
_ 75.6608°W, shrub approx. 10 years old, escaped from plant- 
ings, 27 April 2002, P. M. Catling (DAO sub subsp. mugo). 


In addition to these vouchered observations, 13 sight 
records of young trees spread from cultivation are 
plotted on the accompanying map (Figure 3). 


(5) Identification 

Not all pines are easily identified. The most useful 
texts for identification are Shaw’s (1914) well-illustrated 
compendium and the classic handbook by Dallimore 
et al. (1966). Cope (1986) also provides a key to all of 
the species cultivated in the northeast and a list of cul- 
tivars and their characteristics. Different authors have 
used different characters to distinguish the Asian species, 
and a comprehensive taxonomic study is needed. The 
following provisional key, derived from both previous- 
ly published work and examination of specimens, will 
help to distinguish Black and Mugo Pines from similar 
two-needle pines including some that may have been 
overlooked. 


228 


42° 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


o9P 


oVl 


ov 


oGV 


Pinus mugo (escaped) 


@ herbarium collections 


As sight records 


84° 82° 80° 


= 
(hem 76° 


FIGURE 3. Central and eastern portions of southwestern Ontario showing locations where Mugo Pine (Pinus mugo) has 
escaped from cultivation. Escaped occurrences supported by herbarium specimens at DAO, Agriculture and Agri- 
food Canada, Ottawa, are indicated with a solid dot. Sight records of the author are indicated by solid triangles. 


ta; Reaves'7.5-le Gm long. 3005. cs sneer cote ac 2 
2a Needles snap when bent 180° (at least in P. resinosa); 

cones with or without prickles (recurved 

hook at centre of umbo?) on subterminal scales 

(Figure 4); resin canals marginal*; winter buds more 

of less:neddigh-browat je .0 het hes ey 28 Zi 
3a. Twigs glaucous; cones usually with prickles 

(recurved hook at centre of umbo?) on subterminal 

SOAIESEy itu ack. Pinus densiflora Siebold & Zuccarini, 

JAPANESE RED PINE 

3b. Twigs not glaucous; cones without prickles 

(recurved hook at centre of umbo?) on subterminal 

SCAIESAD 12 Ree Pinus resinosa Ait., RED PINE 
2b. Needles do not snap when bent 180°, but simply 

fold and either crease and remain somewhat folded, 

or regain their original straight appearance upon 

release; cones with prickles (recurved hook at 

centre of umbo’) on subterminal scales (Figure 4); 

resin canals median’; winter buds pale silvery ...... 4. 
4a. Seed cones sessile with rounded base; terminal bud 

resinous; basal leaf sheath ending in a single elongated 

tip; scales of winter buds reddish-brown with white 

CUR eee ame i an ae Pinus nigra Arnold, BLACK PINE 


4b. Seed cones with stalks and truncate at the base; 
terminal bud not resinous; basal leaf sheath 
ending in two long filaments; scales of winter 
buds white Ol) +. 2 ase eee Pinus thunbergii Parlatore, 
JAPANESE BLACK PINE 
lb. Leaves relatively short, 2-7.5 cm long 
5a. Resin canals median? (Figure 5) 
Ie 9, arta. chaweneacdt eee Pinus nigra Arnold, BLACK PINE 
5b. Resin canals marginal or submarginal* (Figure 5)... 6. 
6a. Needles blue-green, often twisted; cones straight; 
upper bark orange-brown; twigs pale yellowish 
Of Greenish 7. . ake: Pinus sylvestris L., SCOTS PINE 
6b. Needles green, not twisted; cones straight or curved; 
upper bark brown; twigs dark brown or greenish .... 7. 
7a. Needles twisted; cones curved 
phil vie ete ame P. banksiana Lamb., JACK PINE 
7b. Needles not twisted; cones, straight ..< <2:%cr n/a 8. 
8a. Leaf margins long-tapered and pointed at the apex; 
leaf sheaths early deciduous leaving pale leaf bases 
on older branches; seeds not winged 
Met ONG Se seats ae Renee P. edulis Engelmann, NuT PINE 
8b. Leaf margins abruptly tapered and somewhat 
rounded at the apex; leaf sheaths persistent and 
therefore bases not pale; seeds winged 


2005 


9a. Cones 2-5 cm long, symmetrical at the base; 
apophysis! flat or slightly elevated but not recurved 
and hooked (Figure 6) ..... P. mugo Turra subsp. mugo 
DwarF Muco (MOUNTAIN) PINE 
9b. Cones 2.5-7 cm long, oblique at the base; 
apophysis prominently recurved and hooded 


Or nooken (ripure'®) i) Wie. ie ing oe oh oben © 10. 
10a. Apophysis! on basal part of outer side of cone 
hooked and recurved ... . PR. mugo Turra subsp. uncinata 


(Ramond) Domin, Swiss MUGO (MOUNTAIN) PINE 

10b. Apophysis! rounded and hooded ..... P. mugo Turra 
nothosubsp. rotundata (Link) Janchen & Neumayer, 
HyBRID MuGo (MOUNTAIN) PINE 


' The apophysis is the part of the seed scale that is exposed in a mature 
closed cone. 

* The umbo is a protuberance on the exposed part of the scale (in an 
unopened cone) representing the apex of the growth of the first 
year. Cones of most pines take two years to mature. 

3 in middle of mesophyll between hypodermis and endodermis. 

4 outer edge of mesophyll adjacent to epidermis and hypodermis. 


Pinus nigra 


Pinus resinosa 


1 cm 


FiGuRE 4. Middle cone scales of pines. Above, Pinus nigra 
cone showing terminal prickle on the umbo which is 
the protuberance on the scale representing the apex 
of first year growth, in this case elevated and pyrami- 
dal (Kleinfeld 1774, DAO). Below, Pinus resinosa 
cone lacking a prickle and with the umbo flattened 
except for small points on its lateral edges (Marcoux 
s. n., DAO 40745). Photos by P. M. Catling. 


CATLING: BLACK PINE AND MUGO PINE IN ONTARIO 


PP 


tEX OE aT Sesser gh, Bintan coat Sa wf 
lint SN 


peck 


Pinus sylvestris 


1mm 


FIGURE 5. Diagrammatic cross sections of pine needles show- 
ing position of resin canals (blackened). Above, Pinus 
nigra with median resin canals. Below, Pinus sylvestris 
with resin canals submarginal and very near to or touch- 
ing the epidermal tissue. Redrawn by P. M. Catling from 
Shaw (1914). 


(6) Additional notes on identification and occurrence 
1. Pinus mugo ssp. mugo, ssp. uncinata and nothossp. 
rotundata (P. montana Miller) 

Pinus uncinata Miller ex Mirbel is sometimes 
placed in synonymy with Pinus mugo Turra, or treated 
as a variety (Pinus mugo var. rostrata Hoopes), but it 
has most recently been treated as a subspecies. It grows 
as a tree to 25 m tall with cones 5-7 cm long. The tree 
(ssp. uncinata) and shrub (ssp. mugo) have hybridized 
extensively (Gaussen et al. 1964) and the hybrids have 
been referred to P. mugo nothossp. rotundata (Link) 
Janchen & Neumayer. Apart from forms only the three 
infraspecific taxa (mugo, uncinata and rotundata) are 
recognized in Christensen’s (1987) classification which 
reduced P. mugo from 16 species with 91 varieties. 

The cones of P. mugo are essentially sessile whereas 
those of P. sylvestris are stalked. Pinus mugo and Pinus 
sylvestris hybridize in their native range to produce 
Pinus xrhaetica Briigger (Christensen 1987). Since 
both occur together as escapes along roadsides in 
Ontario, there is a possibility for this hybrid to occur 
here as well. It possesses characters of P. sylvestris 


230 


Pinus mugo 
ssp. uncinata 


Pinus mugo 
ssp. Mugo 


4cm 


1em 


FIGURE 6. Cones and cone scales of Pinus mugo showing 
apophyses (exposed portion of seed scale). Above, ssp. 
uncinata cone (left) and lateral view of scale (right) 
showing hooked and recurved apophyses (Catling 
2001-11, DAO). Below, ssp. mugo cone (left) and lat- 
eral view of scale (right) showing apophyses elevated 
but not recurved and hooked (Catling 2002-1, DAO). 
Photos by P. M. Catling. 


including the peeling bark, but the bark is more greyish- 
brown than orange and the leaves may be either bright 
green as in P. mugo, or glaucous green as in P. sylvestris. 
As in P. mugo the umbo is bordered by a black, grey 
or dark brown ring (Christensen 1987). 


2. Pinus nigra 

Pinus nigra has been more divided into infrataxa 
(e.g., Bailey 1948; Vidakovic 1974) than other pines 
with similar levels of variability (Earle 2001*). The 
major pattern of variation in Eurasia involves eastern 
and western groups (e.g., Dallimore et al. 1966; Scalt- 
soyiannes et al. 1994): 


la. Needles stout and rigid, 1.5-2 mm thick 

fs Sy ete, subsp. salzmannii (Dunal) Franco (eastern) 
lb. Needles slender and flexible, 0.8-1.5 mm thick 

OER or OE CLE rote subsp. nigra (western) 


The western ssp. nigra includes 3 varieties (Earle 2001): 
la. Needles 6-14 cm; bark on old trees greyish 

var. nigra, AUSTRIAN PINE 

lb. Needles 8-18 cm; bark on old trees orange-pinkish 
2a. Cones yellowish, 5-10 cm long ... var. caramanica 
(Loudon) Rehder, TURKISH PINE 
2b. Cones greyish, 7-12 cmlong ...... var. pallasiana 
(Lambert) Aech. & Graebn., CRIMEAN PINE 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Plants of Pinus nigra that are 1-4 years old have 
needles that are much shorter, often 4-6 cm long, than 
those of older plants, which are usually over 7.5 cm 
long. Consequently P. nigra appears twice in the pre- 
ceding key to species where the first couplet distin- 
guishes species on the basis of needle length. Among 
the few and rather rare dwarf forms of P. nigra is cv. 
hornibrookiana, a shrubby plant with stiff lustrous dark 
needles less than 6 cm long (illustrated by Bailey 
1948, plate IX). The median resin canals readily sepa- 
rate the latter cultivar or a young short-needled plant 
of P. nigra from P. mugo. 

The needles of the Black Pine group that do not snap 
when bent and the presence of spines on the cone scales 
readily distinguish it from Red Pine with which it is 
most likely to be confused. With respect to its separa- 
tion from P. thunbergii, different authors have empha- 
sized different characters. Earle (2001*) suggests that 
the tomentum on the young elongating shoots is dis- 
tinctive in P. thunbergii, but Dallimore et al. (1966) 
describe the young shoots as glabrous. Young plants 
of Black Pine with relatively shorter needles can still 
be distinguished from other short-needled species by 
the position of the resin canals as indicated in the key. 
In the eastern part of their natural range P. nigra and its 
relatives (including P. thunbergii) require more taxo- 
nomic study. Pinus ponderosa Douglas (Ponderosa 
Pine) is sometimes sold and planted in Ontario as P. 
nigra, but the former is readily distinguished by leaves 
usually in fascicles of 3 and by the much more promi- 
nent prickles (1-2 mm in length) on the middle and 
upper cone scales. 

Natural hybridization between Black and Japanese 
Red pines has been reported in southern Michigan 
(Wright et al. 1969). The hybrids were reported to grow 
more rapidly than either parent. At age 4 years they 
could be identified by needles of intermediate stiff- 
ness and sharpness (between the stiff, sharp needles 
of P. nigra and the flexible, more blunt needles of P. 
densiflora). Presence of 3-needled fascicles was also 
characteristic of the hybrid. The young hybrid trees 
are also identifiable by their terminal buds which are 
darker brown than in Black Pine, and by intermediacy 
in position of resin ducts. The Black-Japanese Red pine 
hybrid is considered commercially valuable for both 
timber and roadside planting. In Europe natural hybrids 
have formed with P. mugo and P. sylvestris (e.g., Vida- 
kovic 1958). Many other hybrids have been produced 
to create superior trees for lumber production (van 
Haverbeke 2002*). 

Trees of Pinus nigra reach 6-10 m in height and 
10-15 cm dbh when 15-20 years of age and many pro- 
duce seed as early as age 15. They are reported to have 
growth rates similar to that of the more widely planted 
and native P. resinosa on some sites (van Haverbeke 
2002*). They are not necessarily superior to P. resinosa 
outside of the special circumstances of roadsides (Mor- 
row 1975). Pinus nigra has been recommended as an 


2005 


alternative to P. resinosa, where the latter is particularly 
subject to European pests (Wright and Bull 1962). 


3. Pinus edulis 

Also treated as Pinus cembroides var. edulis, this 
taxon is culitivated for its edible seeds often called 
“pinons”’. It is widely cultivated outside its native range 
which is the semi-desert of the southeastern United 
States. The native range includes climatic zones similar 
to those of southern Ontario. It is occasionally intro- 
duced into remote natural settings such as the Kaladar 
Jack Pine barrens in Ontario (approximately 44.5333°N, 
77.1500°W). Trees begin to bear cones when 25 years 
old and 3 m tall. 


4. Pinus resinosa 

Red Pine is native to southeastern Canada and the 
adjacent United States. It is one of the most widely 
planted pines in Ontario, as individual trees, in small 
plots and in plantations. It spreads readily from plant- 
ings, but to a much lesser extent than Pinus sylvestris. 
In addition to the key characters, the reddish flaky 
bark 1s distinctive. 


5. Pinus sylvestris 

Scots Pine, also called Scotch Pine, is native to Eura- 
sia. It frequently spreads from cultivation in Ontario 
forming dense stands that exclude native species. The 
yellowish or light brown branches and glaucous nee- 
dles of young trees are distinctive. The orange bark of 
the upper trunk and outer branches are distinctive in 
older trees. 


6. Pinus densiflora 

A tree to 40 m tall, P. densiflora is native to Japan, 
Korea, China and Russia. It is very similar to P. syl- 
vestris from which it can be distinguished by longer, 
dull green (but not glaucous) leaves, glabrous branch- 
lets and larger cones. The conelets of P. densiflora are 
erect instead of reflexed as in P. sylvestris (Shaw 1914). 
The 2-4-year-old branches are without exfoliating 
scales unlike those of P. nigra, P. resinosa and P. thun- 
bergii (Bailey 1948). Pinus densiflora is now widely 
planted in Europe and North America. 


7. Pinus thunbergii 

A tree to 43 m tall, P. thunbergii is native to Japan 
and southern Korea. Its pale and rigid leaves are use- 
ful in identification. The fresh cones are brown in P. 
thunbergii instead of brownish-yellow as in P. nigra 
(Shaw 1914). Pinus thunbergii also has fewer and larger 
cone scales than P. nigra, but is apparently closely 
related to the latter species. It has also been separated 
from P. nigra by its tendency to have yellow or orange 
twigs (as in P. resinosa), instead of brown or dark grey 
twigs (e.g., Bailey 1948). 


8. Pinus banksiana 

A tree to 30 m tall, P. banksiana 1s native to the bore- 
al and mixed forest regions of Canada excepting the 
_ western cordillera. It is readily distinguished by crooked 
branches, relatively short needles and curved cones. It 


CATLING: BLACK PINE AND MUGO PINE IN ONTARIO 


231 


has been frequently established in plantations on dry 
sites. 
(7) Prospects 

At the present time neither Black or Mugo pines are 
seriously affecting native biodiversity. Naturally estab- 
lished individuals often exist in small numbers and 
mostly along roadsides and in vegetation comprised 
of other alien species. Although evidently less aggres- 
sive than the introduced Scots Pine (Pinus sylvestris), 
both Black and Mugo pines have a potential for nega- 
tive impact on biodiversity in dry, rocky or sandy habi- 
tats, especially in connection with extensive plantings. 
Black Pine in particular has been shown to grow better 
in North America than in its native range, to reduce 
native biodiversity and cover, and requires relatively 
high levels of herbicide application to control. As a 
result of its superficial similarity to native species, it may 
be overlooked as a problem, thus leading to manage- 
ment difficulties that would not exist if it was identi- 
fied as a risk at an early stage. 


Acknowledgments 

Peter Uhlig and Mike Oldham helped in gathering 
information on the status of pines in Ontario and their 
identification. W. J. Cody and J. Cayouette provided 
useful comments on the manuscript. G. Mitrow assisted 
in obtaining data. 


Documents Cited (marked * in the text) 

Earle, C. J. 2001. Gymnosperm Database: Pinus thunbergii 
Parlatorre. http:/www.conifers.org/pi/pin/th.htm 

van Haverbeke , D. F. 2002. Pinus nigra Arnold, European 
Black Pine. Silvics of North America. United States Depart- 
ment Agriculture. http://www.na.fs.fed.us 

New Zealand Department of Conservation. 2002. South 
Island Wilding Conifer Strategy. http://www.doc.govt.nz/ 
conservation/003~weeds/south-island-wilding-conifer- 
strategy/005~implementing-the-strategy/6.4-control.asp 


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Received 7 January 2003 
Accepted 26 April 2005 


See 


A Potential for the Use of Dragonfly (Odonata) Diversity as a 
Bioindicator of the Efficiency of Sewage Lagoons 


PAUL M. CATLING 
Biology, University of Ottawa, 30 Marie Curie, Ottawa, Ontario KIN 6N5 Canada; catlingp@em.agr.ca 


Catling, Paul M. 2005. A potential for the use of dragonfly (Odonata) diversity as a bioindicator of the efficiency of sewage 
lagoons. Canadian Field-Naturalist 119(2): 233-236. 


In order to determine whether a relationship existed between water quality and odonate fauna in sewage ponds, data were 
gathered at each of six ponds of similar construction and equal size and depth in an adjacent series of improving water quality 
at a lagoon system near Embrun in eastern Ontario. Numbers of nymphs of different species of Odonata were recorded in 
spring and fall, and similar data was collected on adults in June and July. The data on species presence and abundance for 
each of three pairs of cells in the sequence was then compared with the corresponding chemical data which included biological 
oxygen demand, total phophorus, total nitrogen and suspended solids. Water quality improved through the system and 
species diversity in the final ponds was twice that of the ponds receiving wastewater. Numbers of individuals also increased 
through the system. Occurrence of Anax junius, Enallagma civile and Ischnura verticalis alone was associated with poorer 
water quality. Higher diversity including Lestes disjunctus, Leucorrhinia spp. and Erythemis simplicicollis , indicates higher 
water quality. A potential exists for Odonata species diversity, numbers of individuals and occurrence of particular species to 
be used as a bioindicator of water quality and a means of evaluating efficiency of a lagoon system. Advantages include data 


that reflects a time period rather than a point in time and also low costs. 


Key Words: Odonata, dragonflies, bioindicator, biodiversity, sewage lagoons, pollution, water quality, Ontario, Canada. 


Sewage lagoons, a cost-effective form of waste treat- 
ment, are widespread in Canada, especially in sparsely 
populated regions. Many of the present lagoon systems 
in eastern Canada were developed after 1950 (Mano- 
haran 1985). They soon became recognized as a distinc- 
tive environment resembling western prairie sloughs 
in their high productivity and biological composition. 
Lagoon systems are characterized by a series of pools 
of increasing water quality from the first receiving 
water with raw sewage to the last receiving water that 
has had substantial treatment. This sequence provides 
an opportunity for assessing association of aquatic 
species and abundance in relation to water quality. 
Such information can be applied to an assessment of 
the efficiency of a lagoon system. 

Biodiversity and numbers of individuals of dragon- 
flies have been shown to be related to water quality 
worldwide (e.g., Watson et al. 1982; Takamura 1991; 
Corbet 1999) and in Canada (Lefort and Catling 1998), 
but their potential use in evaluating sewage lagoon 
systems has not been studied. A preliminary investiga- 
tion (Catling 2004) suggested a relationship between 
water quality and dragonfly species presence and abun- 
dance in sewage lagoons and identified species groups 
representative of good and poor water quality. Although 
useful, this study was based indirectly on water quality 
through a demonstrated correlation with water clarity. 
The present work adds to this by elucidating the direct 
relationship with specific values of water quality param- 
eters in a series of sewage ponds. The objective was to 
further explore the extent to which dragonflies have a 
potential use in evaluating the efficiency of sewage 
lagoon systems. 


Methods 

Each of six ponds of similar construction and equal 
size and depth in an adjacent series of improving 
water quality were sampled at a lagoon system near 
Embrun (45.2767°N, 75.2359°W) in eastern Ontario. 
Numbers of nymphs of different species of Odonata 
were recorded in spring and fall (15 October 2001 
and 17 April 2002), and similar data were collected on 
adults in summer (26 June 2002 and 18 July 2002). 
The data on species presence and abundance for each 
of three pairs of cells in the sequence were then com- 
pared with the corresponding chemical data which in- 
cluded biological oxygen demand (BOD), total phopho- 
rus (TP), total nitrogen (TN) and suspended solids (SS). 
The data included three measurements (fall 2001, spring 
2002, fall 2002) of each parameter from each of the 
Six sewage ponds. All measurements (mg/L) were made 
at Accutest Laboratories in Ottawa on behalf of the 
township of Russell. These measurements were aver- 
aged to provide an indication of overall conditions. 
Since adjacent sewage ponds and even parts of ponds 
appear to vary greatly in water chemistry due to the 
time of year, amount of input and other factors relat- 
ing to speed of breakdown, the point samples can only 
be considered to indicate a trend in the sequence. To 
compensate for variation, chemical data in each adja- 
cent pair of cells (in the sequence of six) were averaged. 
This averaging also accounted for the bypassing of cells 
1, 3 and 5 during periods of heavy use, making certain 
sequential pairs of cells similar. 

Nymphs were sampled with 50 scoops of a net at 
each of the six ponds on each of two visits. Thus it was 
the potential and successful overwintering nymph pop- 


233 


234 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |. Number of nymphs of species of Odonata found in spring (S), fall (F) and the total (T) for three pairs of cells at the 


Embrum lagoon system. 


Species 1,2 

S F at 
Anax Junius — 2 2D 
Enallagma civile - 50 50 


Enallagma cyathigerum - - — 
Erythemis simplicicollis ~ - _ 


Ischnura verticalis 10 150 ~—-:160 
Lestes disjunctus - - ~ 
Leucorrhinia proxima C= - - 
Leucorrhinia intacta - - - 
Libellula quadrimaculata ~ 10 10 


Libellula pulchella = = = 


TABLE 2. Total number of adults of species of Odonata recorded on two summer visits to the Embrun lagoons. 


Species 1,2 
Anax junius 5 
Enallagma civile S) 


Enallagma cyathigerum ~ 
Epitheca cynosura = 
Ischnura verticalis 10 
Lestes disjunctus - 
Tramea lacerata - 


ulations that were sampled. Sample points were approx- 
imately equally distributed around the edges of the 
lagoons. The nymphs were identified using the keys in 
Walker (1953). The heavily mottled body, long-pointed 
caudal lamellae and eyes with horizontal black lines, 
made the identification nymphs of Eastern Forktail 
(Ischnura verticalis) relatively simple. Nymphs of the 
species of bluets (Enallagma) were less easily identi- 
fied. They were assigned to taxa in the field based on 
characteristics of the caudal lamellae evaluated with a 
15x hand lens. In cases where the numbers of larvae 
exceeded a hundred, species composition was often 
based on microscopic examination of 20-50% of the 
total identified in the field. Approximately 95% of field 
identifications were correct. Adults were surveyed over 
a period of two hours on each of two dates with approx- 
imately 20 minutes spent at each of the six ponds. 


Results and Discussion 

At the Embrun lagoon system it is clear that bio- 
logical oxygen demand, total nitrogen and suspended 
solids decrease from the lagoons receiving wastewater 
to the final “polishing ponds” that release treated efflu- 
ent back to surface water drainage (Figure 1). Total 
Phosphorus is highest in the mid-portion of the system, 
possibly due to gradual release from organic matter 
and then precipitation in the middle part of the process. 
The water in the final pool of the Embrun system is 


3,4 5,6 
S F F S F [ 

1197, 80 ZY 10 35 45 
24 210 234 100 265 365 
= = = 60 a oF 
= = ~ l _ | 
34 135 169 20 160 180 
= = = - 8 8 
= = — 20 ~ 20 
= = = 3 = 3 
= = = a 3) 10 
- - ~ 7 3 10 

3,4 5,6 

30 55) 

400 795 

80 60 

6 3 

190 242 


relatively clean and well below the 25 mg/L compli- 
ance criteria for biological oxygen demand (BOD) and 
suspended solids (SS). Associated with this increase 
in water quality through the series of ponds, both the 
numbers of individuals and the numbers of species of 
dragonflies increases (Tables 1 and 2, Figure 2). Num- 
bers of individuals of nymphs were not that much 
higher in the final ponds than in the middle ponds in 
the series (Figure 2 ), but there were large numbers of 
the few species present. The increase in diversity of 
species and numbers of individuals with improving 
water quality supports similar findings in studies com- 
paring polluted and non-polluted waters (e.g., Watson 
et al. 1982; Takamura 1991; Lefort and Catling 1998; 
Corbet 1999). 

Northern Spreadwing (Lestes disjunctus), whitefaces 
(Leucorrhinia spp.) and Eastern Pondhawk (Erythemis 
simplicicollis) were found only in the ponds with supe- 
rior water quality as was the case in a preliminary study 
of several eastern Ontario lagoon systems based on 
correlates of water quality rather than actual values 
(Catling 2004). In addition the three species charac- 
teristic of the lowest water quality in that correlative 
study were same three found alone in the poor quality 
ponds at Embrun; i.e., Common Green Darner (Anax 
Junius), Familiar Bluet (Enallagma civile) and Eastern 
Forktail (/schnura verticalis). Data are insufficient to 
give an accurate indication of the tolerance limits of 


2005 
24 
BIOLOGICAL 
28 OXYGEN 
DEMAND 
16 
l2 
8 
4 
oe ee’) 
Te) 
S te 3,4 
18 
TOTAL 
9 NITROGEN 


1,2 3,4 5,6 


CATLING: DRAGONFLY DENSITY AT A BIOINDICATOR 


235 


SUSPENDED 
SOLIDS 


TOTAL 
PHOSPHOROUS 


1,2 3,4 5,6 


Treatment ponds (in sequence) 


different species, but average chemical conditions asso- 
ciated with dragonfly populations can be estimated 
from Figures | and 2. 

The first lagoons were poorer in species diversity and 
numbers in the spring than in the fall possibly due to 
decreased rate of decomposition and reduced oxygen 
during the winter months resulting in extreme condi- 
tions that eliminated much of the fauna in the initial 
ponds. Nymphs in early spring may provide the best 
indication of water quality due to direct association with 
water following a period of stress. However, adults ap- 
pear to be able to recognize water quality and some 
species tend to occur only on the cleaner ponds (Fig- 
ure 2). Overall the increase in species diversity in the 
final ponds is twice that of the ponds receiving waste- 
water. 


FIGURE |. Mean measurements of chemical parameters at Embrun, eastern Ontario, based on sampling of the six ponds on 
three separate dates (22 October 2001, 26 February 2002, 15 October 2002). 


This study suggests that the improving water quality 
in a series of sewage ponds is reflected by increasing 
odonate species diversity and increasing numbers of 
individuals of both aquatic larvae and aerial adults. 
The use of the indicator species is particularly promis- 
ing and can be used in conjunction with diversity and 
numbers. The potential to use dragonflies as a bioindi- 
cator of the efficiency of sewage ponds requires addi- 
tional study to confirm this suggestion over a broad 
area. It has two obvious advantages over chemical 
tests: (1) It includes reference to a time period (the 
larvae living in the pond for at least several weeks) 
rather than a single chemical sample at one particular 
point in time that may or may not be representative 
of conditions over the longer period. (2) It is inex- 
pensive and can be done at most times of year using 


236 


18 


SPECIES 
OF ADULTS 


120@ 


NUMBER 
OF ADULTS 


1828 


808 


680 


498 


1,2 3,4 5,6 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


18 


SPECIES 
OF NYMPHS 


@ 
1,2 3,4 5,6 
920 
NUMBER 
800 OF NYMPHS 


1,2 3,4 5,6 


Treatment ponds (in sequence) 


FIGURE 2. Numbers of species and individuals of both larval and adult Odonata a the Embrun lagoons. 


either mature larvae or adults, or both depending on 
the time. 


Acknowledgments 

Craig Cullen, superintendent of public utilities for 
the township of Russell, assisted by providing exten- 
sive information on the lagoon systems at Embrun. 


Literature Cited 

Catling, P. M. 2004. A preliminary study of dragonflies at 
eastern Ontario sewage lagoons in relation to water quality. 
Pages 28 — 32 in Ontario Odonata, volume 4. Edited by P. 
M. Catling, C. D. Jones, and P. Pratt. Toronto Entomolo- 
gists’ Association, Toronto, Ontario, Canada. 

Corbet, P. S. 1999. Dragonflies, behaviour and ecology of 
Odonata. Cornell University Press, Ithaca, New York. 829 
pages. 


Lefort, F., and P. M. Catling. 1998. A survey of damselfly 
adults at urban and non-urban streams at Ottawa, Ontario. 
Argia — the news journal of the dragonfly society of the 
Americas 10(4): 17-19. 

Manoharan, R. 1985. Sewage lagoons in cold climates. 
Environmental Protection Service (Environment Canada) 
EPS4/NR/1. 89 pages. 

Takamura, K., S. Hatakeyama, and H. Shiraishi. 1991. 
Odonate larvae as an indicator of pesticide contamina- 
tion. Applied Entomology and Zoology 26: 321-326. 

Walker, E. M. 1953. The Odonata of Canada and Alaska, 
volume |, part 1: General. part 2: The Zygoptera — Dam- 
selflies. University of Toronto Press. 292 pages. 

Watson, J. A. L., A. H. Arthington, and D. L. Conrick. 
1982. Effect of sewage effluent on dragonflies (Odonata) 
of Bulimba Creek, Brisbane. Australian Journal of Fresh- 
water Research 33: 517-528. 


Received 30 April 2003 
Accepted 26 April 2005 


Invasive Scots Pine, Pinus sylvestris, Replacing Corema, Corema 
conradii, Heathland in the Annapolis Valley, Nova Scotia 


PAUL M. CATLING! and SUSAN CARBYN2 


Agriculture and AgriFood Canada, Environmental Health, Biodiversity, William Saunders Building, Central Experimental 
Farm, Ottawa, Ontario KIA 0C6 Canada; email: catlingp @agr.gc.ca 

Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, 32 Main Street, Kentville, Nova Scotia B4N 1J5, 
Canada 


Catling, Paul M., and Susan Carbyn. 2005. Invasive Scots Pine (Pinus sylvestris) replacing Corema, Corema conradii, heathland 
in the Annapolis valley, Nova Scotia, Canadian Field-Naturalist 119(2): 237-244. 


Examination of air photos from 1930, 1970 and 2002 revealed stands of the European Scots Pine (Pinus sylvestris) invading 
remnants of natural Corema (Corema conradii) heathland in the Annapolis valley. To document the impact of the introduced 
pines, four natural habitats were compared with two adjacent habitats already invaded by the pines. All surveyed habitats 
had been dominated by Corema heath based on air photos taken in 1930. Twenty | m? quadrats were used to record pres- 
ence and cover of vascular plants at each site. The invasive alien pines reduce the native cover to 12%. Vascular plant biodi- 
versity is reduced to less than 42% and the cover of the heathland dominant, Corema conradii, is reduced from over 100 % to 
less than 2%. with Deschampsia flexuosa becoming the dominant species. The modified ecosystem and loss of biodiversity has 
economic impacts through loss of pollinators of agricultural crops and loss of germplasm of native crop relatives. 


Key Words: Scots Pine, Pinus sylvestris, Corema, Corema conradii, invasive, alien, heathland, barrens, Nova Scotia, Canada. 


Scots Pine (Pinus sylvestris L.) 1s native to much 
of Europe and northern Asia and has been introduced 
and become naturalized throughout the temperate 
regions of the world. It is the most widely distributed 
of the world’s pines (e.g., Farrar 1995) and is used 
for landscaping, shelterbelts, soil stabilization and 
general reforestry purposes. Not only is it used for all 
of these purposes in Canada but is also cultivated 
extensively for use as Christmas trees. In some parts 
of Canada it has spread from cultivation and invaded 
natural habitats (Catling 1997). 

One of the places where a significantly deleterious 
invasion is occurring is in the remnants of the heath- 
land ecosystem dominated by Corema (Corema con- 
radii Torr.) on the sand plains of the Annapolis valley, 
Nova Scotia. These threatened plant communities are 
already deteriorating due to succession of the vegetation 
to forest apparently as a result of the lack of fires. Mesic 
sites are the most susceptible. Although native woody 
vegetation is slow to invade the drier sites, Scots Pine 
has proved to be a very aggressive invader that forms 
dense stands that exclude most other plant species. 

Here we document the conversion of natural heath- 
| land to Scots Pine wood and provide an indication of the 
| vegetation changes that have likely occurred through 
/a comparison of invaded and non-invaded sites. This 
will permit a better understanding of vegetation dynam- 
| ics in the heathland and will provide the information 
needed for effective management. 


| Methods 

Documenting invasion of Scots Pine 

Through examination of air photos from 1930, 
; 1970 and 2002 aerial surveys (Figure 1) and exami- 


nation of sites in 2003, specific areas of heathland 
that have been invaded by Scots Pine can be clearly 
identified providing an opportunity to assess the impact 
of the invading pines. These areas have evidently not 
been subject to other unnatural disturbances so that 
the differences in vegetation between them and the 
adjacent heathland are due to Scots Pine invasion. The 
stands of Scots Pine on the heathland appear to have 
originated from seed dispersed from trees planted 
around buildings or in plantations nearby. Isolated older 
Scots Pines 10-30 years of age produce seed readily 
and are frequently surrounded by younger trees 1- 15 
years old which grow with further recruitment to form 
a dense stand within another 15-20 years. 


Assessing impact of Scots Pine 

The conversion of natural heathland to Scots Pine and 
specifically the impact of Scots Pine was assessed 
through comparison of invaded heathland with adja- 
cent natural heathland with various amounts of native 
tree cover. The study area was located at approx. 
45.0166°N, -64.8833°W between Auburn and Kingston 
in Annapolis County, Nova Scotia. On 26 June 2003, 
presence and cover of vascular plant species was recorded 
in six habitats within an area that was essentially 
open heathland 50 years earlier based on aerial photo- 
graphs on file at Nova Scotia Department of Natural 
Resources (Figure 1). Of the six habitats sampled 
four were dominated by native species of open heath- 
land and two were dominated by Scots Pine which 
had invaded parts of the heathland. The two open 
heathland sites described were immediately adjacent to 
the Scots Pine stands and the introduced pines were 
extending into the open heathland area. The two heath- 
land sites with Red Pine (Pinus resinosa Ait.) had 


257 


238 


é 


FiGurE |. Airphotos showing a portion of the heathland on the west side of Auburn (approx. 45.0166° N, - 64.8833° W) in 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


1930, 1970 and 2002. The stands of Pinus sylvestris where ground flora was evaluated are delineated with a black 
line on the 2002 photo. In 1930 heathland existed where these stands now occur. The Scots Pine stands were just 


starting to develop in 1970. 


adjacent open heathland. All sites sampled were evi- 
dently Corema-dominated heathland in 1930 (Figure 1). 

Site 1, an open heathland approx. 90% open in aerial 
view with scattered White (Pinus strobus L.), Red, and 
Scots pine and site 5, the adjacent Scots Pine stand was 
located at 45.0107°N, 64.8960°W. Site 2, the other open 
heathland, also approx. 90% open, and site 6, the 
adjacent Scots Pine stand was located at 45.0157°N, 
-64.8803°W. Site 3, an open heathland, 60% open in 
aerial view, with Red Pine 50 years old was located at 
45.0014°N, -64.9419°W. Site 4, the semi-open 60-70 


year-old Red Pine stand, approximately 20% open in 
aerial view was located at approximately 44.9916°N, 
64.9750°W. 

At each site the habitat was delineated and within it a 
transect was selected at random using a compass line 
derived from random numbers. Presence and cover was 
recorded for each species in each of 20 one m? quadrats 
at 5 m intervals along the transect line. Cover was esti- 
mated as 2 the surface area of each species rather than 
a simple aerial view and thus could exceed 100% of 
the surface of the square metre sampled. This method 


2005 


CATLING AND CARBYN: SCOTS PINE REPLACING COREMA HEATHLAND 


239 


FiGurE 2. Natural heathland on the west side of Auburn (approximately 45.0166° N, 64.8833° W) with ground cover domi- 


nated by Corema (Corema conradii) and scattered native Red Pine (Pinus resinosa). 


provided a maximally accurate description of biomass. 
Species up to 2 m tall (including trees) were recorded. 
In a very few cases where a large tree trunk was in a 
selected quadrat, the position of the quadrat was moved 
(up to 2 m) to avoid the trunk. For comparison, the 
mean frequencies and covers for each pair of commu- 
nities was calculated and expressed as a percentage 
of that of the open heath. Voucher specimens are in 
the collection of Agriculture and Agri-Food Canada 
in Ottawa (acronym DAO). 


Results and Discussion 
Native Red Pine and White Pine exist as scattered 
trees in open heathland (Figure 2) and the mature trees 
are rarely accompanied by nearby saplings and seed- 
lings, presumably due to difficulty of establishment and 
specific requirements such as suitable mineral substrate. 
| In contrast mature Scots Pines are usually surrounded 
| by saplings and seedlings, many of which survive to 
form a dense stand, and the trees can establish in organ- 
| ic soils. 
Red Pine forming stands in open heath (Figure 3) 
May result in a reduction of low heathland cover to 
47.5%, as well as a reduction of diversity to 75.7% 


(Tables | and 2). The dense stands of Scots Pine that 
develop can cause a much greater reduction of cover 
and biodiversity with cover reduced to 12.0% and diver- 
sity reduced to 42.4% (Figures. 4 and 5). Corema, the 
dominant of open heath is reduced from over 100 % 
cover to less than 2 % cover in stands of Scots Pine 
(Figure 5). Other than Scots Pine, invasive alien species 
were not prominent in the sites studied and they did not 
increase in the presence of Scots Pine (Tables | and 2). 

After invasion by Scots Pine forest, Deschampsia 
flexuosa replaces Corema conradii as the dominant vas- 
cular plant and ground cover is greatly reduced (Tables 
1 and 2). Significant species, including Amelanchier 
lucida Fernald, Vaccinium angustifolium Ait., and 
species of Rubus, all of which provide food for pollina- 
tors that service local crops (apples, blueberries, etc) 
are severely reduced after invasion by Scots Pine. 
These same species and several others are also poten- 
tially important as sources of genetic variation for crop 
improvement. 

The numbers of immature Scots Pines in open heath- 
land ranged from 80—350 per acre and were 99 % of the 
trees on open heath adjacent to the Scots Pine stands. 
Native pines and other trees apparently invade the drier 


240 THE CANADIAN FIELD-NATURALIST Vol. 119 


TABLE |. Frequency of species less than 2 m high in three major associations, open heath, Red Pine heath and Scots Pine 
forest. The recording was done in 2003 in an area that was a natural heathland dominated by Corema with Red and White | 
Pines 50 years earlier. Introduced species are indicated with an asterisk (*) following the species name. | 


Species Open Heath Red Pine Heath Scots Pine 
1 2 3 4 5 6 

Acer rubrum L. - - 10 ~ ~ 5 
Amelanchier lucida Fern. pa) 35 5 - ~ 5 
Amelanchier laevis Wieg. - 5 5 ~ ~ - 
Arctostaphylos uva—ursi (L.) Spreng. 20 15 10 25 ~ 
Aronia melanocarpa (Michx.) Ell. 30 - ~ - ~ _ 
Betula populifolia Marshall 10 - ~ - ~ _ 
Carex tonsa (Fern.) Bickn. var. 

rugosperma (Mack.) Crins 15 — 5 ~ - ~ 
Comptonia peregrina (L.) Coult. 15 20 2) 5 ~ - 
Corema conradii Torr. 95 95 60 35 30 - 
Cypripedium acaule Aiton ~ - 20 25 ~ ~ 
Danthonia spicata (L.) Beauv. = 25 — — _ — 
Deschampsia flexuosa (L.) Trin. 20 35 oo 50 65 7S 
Dichanthelium acuminatum (Sw.) Gould & Clark 

ssp. fasciculatum (Torr.) Freckman & Lelong 5 — — - — - 
Dichanthelium depauperatum (Muhl.) Gould 5 - - — - - 
Festuca filiformis Pourret * - 30 - ~ ~ - 
Gaultheria procumbens L. 5 a _ ~ — 5 
Gaylussaccia baccata (Wang.) K. Koch = _ — 10 — = 
Hieracium pilosella L. * - - 5 - = 10 
Hudsonia ericoides L. 10 - - - ~ ~ 
Hypericum perforatum L. * | ~ - ~ - - 
Juniperus communis L. 5 = — ~ ~ 5 
Kalmia angustifolia L. 10 20 - - - - 
Maianthemum canadense Desf. - - 2) - - 3 
Melampyrum lineare Dest. ~ 20 30 20 3 ~ 
Monotropa hypopithys L. - - - - 5 - 
Pinus resinosa Att. — - ~ 5 - - 
Pinus strobus L. - 15 - - - 
Pinus sylvestris L. * 15 15 - - 5 - 
Pteridium aquilinum (L.) Kuhn. - 10 - - ~ — 
Quercus rubra L. - - 5 - - ~ 
Rubus vermontanus Blanch. a) =) _ ~ ~ - 
Rumex acetosella L. * 10 ~ _ - - - 
Solidago bicolor L. i 35 5 - 10 - 
Solidago juncea Ait. - >) - - - - 
Trientalis borealis Raf. — — 35 - ~ J 
Vaccinium angustifolium Ait. 50 70 90 80 20 10 
Viburnum nudum L. var. cassinoides (L.) Torr. & Gray _ 5 _ — - - 
Total native species 17 16 16 5 6 8 
Total introduced species | 2 l 0 | l 
Total species 20 18 iL 9 Hi | 
Mean community frequency 16.5 123 7 


% of mean open heath frequency 100.0 Hoe 42.4 


2005 CATLING AND CARBYN: SCOTS PINE REPLACING COREMA HEATHLAND 241 


TABLE 2. Mean cover of species less than 2 m high in three major associations, open heath, Red Pine heath and Scots Pine 
forest. The recording was done in 2003 in an area that was a natural heathland dominated by Corema with Red and White 
Pines 50 years earlier. Introduced species are indicated with an asterisk (*) following the species name. 


Species Open Heath Red Pine Heath Scots Pine 
l 2 3 4 5 6 

Acer rubrum L. — ~ 0.10 - - 0.45 
Amelanchier lucida Fern. 4.80 TEE 0.05 ~ - 0.50 
Amelanchier laevis Wieg. - 2.00 0.15 - - - 
Arctostaphylos uva-ursi (L.) Spreng. 1.60 11.50 LS wpe ~ - 
Aronia melanocarpa (Michx.) Ell. 3:95 - - - - - 
Betula populifolia Marshall 2:55 — — ~ - - 
Carex tonsa (Fern.) Bickn.var. rugosperma 

(Mack.) Crins 0.50 _ 0.05 _ _ - 
Comptonia peregrina (L.) Coult. 4.75 1.85 1:25 0.25 ~ _ 
Corema conradii Torr. 105.00 110.25 13.65 6.10 130 - 
Cypripedium acaule Aiton - ~ 0.35 0.60 ~ - 
Danthonia spicata (L.) Beauv. - 0.25 _ _ — ~ 
Deschampsia flexuosa (L.) Trin. 0.50 1.50 11.10 14.10 17.65 13.40 
Dichanthelium acuminatum (Sw.) Gould & Clark 

ssp. fasciculatum (Torr.) Freckman & Lelong 0.05 ~_ — = _ = 
Dichanthelium depauperatum (Muhl.) Gould 0.05 ~ - ~ - - 
Festuca filiformis Pourret * - 0.30 — - — - 
Gaultheria procumbens L. 1.50 0.20 _ - — 0.40 
Gaylussaccia baccata (Wang.) K. Koch - _ _ 2.85 _ - 
Hieracium pilosella L. * - - 0.10 = - 0.45 
Hudsonia ericoides L. 0.75 - - - - ~ 
Hypericum perforatum L. * 0.12 - ~ - ~ - 
Juniperus communis L. 2.50 - - ~ - 0.35 
Kalmia angustifolia L. 5.90 2.95 ~ ~ - - 
Maianthemum canadense Desf. - - 11.80 ~ - 0.35 
Melampyrum lineare Dest. - 0.80 0.35 5.95 1.00 - 
Monotropa hypopithys L. - - - - 0.10 - 
Pinus resinosa Ait. ~ - - 0.05 - - 
Pinus strobus L. - - 0.35 ~ - - 
Pinus sylvestris L. * 3.00 26 - ~ 0.25 = 
Pteridium aquilinum (L.) Kuhn. ~ a - _ = = 

| Quercus rubra L. - - 0.05 ~ - 


Rubus vermontanus Blanch. 0.24 0.60 - - ~ - 
Rumex acetosella L. * 3.20 - 
Solidago bicolor L. O25 2.10 0.05 - 0.50 ~ 
Solidago juncea Ait. 0.10 0.25 — — 
Trientalis borealis Raf. - _ 0.95 - _ 0.05 


Vaccinium angustifolium Ait. 5.65 31.45 295 43.95 2.0) 0.45 
| Viburnum nudum L. var. cassinoides 
| (L.) Torr. & Gray ~ 0.10 - _ ~ = 
| Total for all species 146.26 185.15 71.05 fo i) 23.50 16.40 
Mean community cover 162.89 TI39 19.6 


| % of mean open heath cover 100.0 47.5 12.0 


242 


by blueberry (Vaccinium angustifolium). 


open heathland very slowly or not at all, whereas the 
European Scots Pine is an aggressive invader that forms 
dense stands. These not only reduce the biodiversity 
of native vascular plant species, but they also eliminate 
the native open heathland ecosystem. Management of 
Scots Pine will be necessary to protect representative 
sites. 

The threat of Scots Pine to natural ecosystems is 
apparently best documented in New Zealand where 
“wilding” (escaped from cultivation) conifers have had 
a major impact on the natural landscape. Of these 
conifers, Scots Pine is rated among the species having 
the greatest impact on the natural ecosystem and sub- 
stantial effort has been directed to control and man- 
agement (e.g., Langer 1992). 

In the United States, various agencies have recom- 
mended against the planting of Scots Pine due to its 
invasive tendency, but, as is the case for Canada, 
quantification of impact is very limited. The lack of 
detail is in direct contrast to the abundance of anec- 
dotes. For example it has been noted that: “to appre- 
ciate the problem of invading alien trees and shrubs 
in Canada, one need only consider the planting of Scots 
Pine (Pinus sylvestris) in native open habitats that have 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


become thoroughly choked with this aggressive weed | 
tree” (Catling 1997). Scots Pine was featured as one | 
of 16 invasive aliens of wetland habitats in Canada and | 
one of 40 invasives of upland habitats in Canada (White | 
et al. 1993). It has also been mentioned in the context | 
of serious invasives in various provinces (e.g., Kaiser | 
1983; Riley 1989; Urban Forest Associates Inc. 2002). | 
and environmental organizations have advised against — 
its use as an ornamental. 

The demonstrated impact on a portion of the declin- 
ing Nova Scotian heathland ecosystem is significant | 
as one of the first quantified examples of impact of | 
invasive alien Scots Pine in North America. It should 
serve as a basis for monitoring impacts in Nova Scotia | 
and elsewhere. Clearly the concerns that have been | 
widely expressed by environmental organizations and | 
field biologists for impact of invasive alien Scots Pines 
are justified. 

In addition to reduction of Corema heathland due to | 
succession and invasive Scots Pine, there has also | 
been substantial reduction due to urban and agricultural | 
development (Figure |) and decline of this ecosystem 
has been rapid over the past few decades. The need to | 
protect representative examples is well known. The | 


2005 CATLING AND CARBYN: SCOTS PINE REPLACING COREMA HEATHLAND 243 


TF | 


3 
: 
: 

54 
| 
3 
by | 


Na 


FiGure 4. Scots Pine (Pinus sylvestris) of many different age classes invading open heathland dominated by Corema (Corema 
conradii) on the west side of Auburn (approximately 45.0083° N, 64.8916° W). 


244 


oT, 


(approximately 45.0166° N, 64.8833° W). 


remnants of this ecosystem provide pollinators for 
adjacent fruit crops and contain distinctive variants 
of native crop relatives that could be valuable in crop 
improvement and diversification (see above and Catling 
et al. 2004). Thus they are sources of valuable biodi- 
versity. The perpetuation of this important ecosystem 
will require not only a system of protected sites but 
also management of succession and invasive aliens, 
particularly Scots Pine. 


Acknowledgments 

Frances MacKinnon at Nova Scotia Department of 
Natural Resources assisted with access to air photos. 
Useful comments were provided by W. J. Cody and 
J. Cayouette. Field assistance was provided by S. P. 
Vander Kloet and S. Javorek. 


Documents Cited (marked * in text) 

Urban Forest Associates Inc. 2002. Invasive exotic species 
ranking for southern Ontario. 7 pages. http://www.serontario. 
org/pdfs/exotics.pdf. (Accessed April 2005). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


‘ beg ae f ¥ Le 


FIGURE 5 Dense stand of Scots Pine (Pinus sylvestris) with ground covered mostly by pine litter on the west side of Auburn 


Literature Cited 

Catling, P. M. 1997. The problem of invading alien trees 
and shrubs: some observations in Ontario and a Canadian 
checklist. Canadian Field-Naturalist 111: 338-342. 

Catling, P. M., S. Carbyn, S. P. Vander Kloet, K. MacKen- 
zie, S. Javorek, and M. Grant. 2004. Saving Annapolis 
Heathlands. Canadian Botanical Association Bulletin 37 
(1): 12-14 

Farrar, J. L. 1995. Trees in Canada. Canadian Forest Service, 
Natural Resources Canada. Ottawa. 502 pages. 

Kaiser, J. 1983. Native and exotic plant species in Ontario: 
a numerical synopsis. The Plant Press 1: 25-26. 

Langer, E. R. 1992. Chemical control of wilding conifer 
seedlings in New Zealand. Plant Protection Quarterly 7(3): 
135-139. 

Riley, J. L. 1989. More invasive aliens. Seasons 29(2): 23. 

White D. J., E. Haber, and C. Keddy. 1993. Invasive plants 
of natural habitats in Canada: an integrated review of wet- 
land and upland species and legislation governing their 


control. Canadian Wildlife Service, Ottawa, Canada. 121 — 


pages. www.cws-scf.ec.gc.ca/publications/inv/cont_ e.cfm 


Received | April 2004 
Accepted 18 April 2005 


Insect Visitation to Wildflowers in the Endangered Garry Oak, 
Quercus garryana, Ecosystem of British Columbia 


A. L. PARACHNOWITSCH and E. ELLE 
Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia VSA 1S6 Canada 


Parachnowitsch, A. L., and E. Elle. 2005. Insect visitation to wildflowers in the endangered Garry Oak, Quercus garryana, 
Ecosystem of British Columbia. Canadian Field-Naturalist 1 19(2): 245-253. 


The Garry Oak Ecosystem (GOE) is a fragmented and endangered ecosystem in Canada, and is currently the focus of conserva- 
tion and restoration efforts in British Columbia. However, little is known about the basic biology of GOE forbs, or their rela- 
tionships with pollinating insects. We monitored wildflowers and their insect visitors in 25 quadrats within a 25 x 25 metre 
plot, located in a fragment of the GOE near Duncan, British Columbia, for six weeks (the majority of the flowering period). 
Overall, 21 native and non-native forb species flowered in our quadrats during the survey, and we observed an additional six 
forb species flowering outside of our quadrats. Eight forbs were visited within quadrats by a total of 13 insect taxa, identified 
to morphospecies. Visits by eight additional morphospecies were observed outside of the quadrats. In general, visitation was low; 
however, most insect morphospecies were observed visiting more than one plant species, and most plant species were visited 
by more than one insect morphospecies, suggesting that pollination may be generalised in this community. A 7 analysis 
indicated that insect visitation was not proportional to the relative abundance of forbs, with higher than expected visitation to 
Common camas (Camassia quamash), and no observed visits to 11 species, most with very small (putatively unattractive) flo- 
wers. The most frequent insect visitor was the introduced Honeybee, Apis mellifera, followed by native mason bees (Osmia 
spp.) and mining bees (Andrena spp.). Our observations provide baseline data for future, detailed studies that should investigate 
the importance of plant-pollinator mutualisms for sustainability of populations and communities in this rare ecosystem. 


Key Words: Garry Oak Ecosystem, Quercus garryana plant-pollinator interactions, Common Camas, Camassia quamash, 


British Columbia. 


The Garry Oak Ecosystem (GOE), an open wood- 
land habitat dominated by Garry Oak (Quercus gar- 
ryana), is increasingly of conservation concern. The 
GOE ranges from California to British Columbia 
(Erickson 2000), and within British Columbia is res- 
tricted to dry, nutrient-poor sites within the Coastal 
Douglas-fir biogeoclimatic zone (Klinka et al. 1996). 
Since western settlement, the GOE has become highly 
fragmented, with as little as 1% of its original area still 
remaining (Fuchs 2001). There are currently 117 GOE 
plant and animal species listed as endangered, threat- 
ened or vulnerable by the Committee on the Status of 
Endangered Wildlife in Canada (COSEWIC), and/or 
the British Columbia, Ministry of Sustainable Resource 
Management, and in Canada, the entire ecosystem is 
considered endangered (Garry Oak Recovery Team 
2003*). 

Historically, the GOE has been maintained by fire 
disturbance and large-scale aboriginal cultivation (har- 
vesting, weeding, replanting and controlled burning) 
of Camassia quamash (Common Camas) bulbs (Thil- 
enius 1968; Turner and Kuhnlein 1983; MacDougall et 
al. 2004). However, European settlers imposed fire sup- 
pression and livestock grazing, leading to the encroach- 
ment of conifers into the GOE, and also introduced 
several highly invasive species; e.g., Cytisus scoparius 
(Scotch Broom) and Poa pratensis (Kentucky Blue 
Grass; Thilenius 1968; MacDougall et al. 2004). Fire 
suppression, non-native species introductions, and 
conversion of land to agricultural or urban use have 


fragmented the GOE, and current research aims to 
understand the implications of this fragmentation for 
Q. garryana establishment (Fuchs et al. 2000; Regan 
and Agee 2004) and to manage invasive and native 
species (e.g., Ussery and Krannitz 1998; Tveten and 
Fonda 1999; MacDougall and Turkington 2004). How- 
ever, little is known about the basic ecology of native 
herbaceous forb communities within the GOE beyond 
status reports of a few rare species (Douglas and Illing- 
worth 1997, 1998; Douglas and Ryan 1998; Penny and 
Douglas 2001) and studies of the reproductive biology 
of Aster curtus (currently Sericocarpus rigidus; Clam- 
pitt 1987; Bigger 1999; Giblin and Hamilton 1999), 
One area of special concern is the paucity of data on 
plant-pollinator interactions within the GOE (Fuchs 
2001). Pollination can be central to recruitment and 
maintenance of individual species, and can thereby 
provide an essential ecosystem service (Kearns et al. 
1998; Black et al. 2001). Habitat loss and fragmenta- 
tion, agricultural practices such as herbicide and pes- 
ticide use, and the encroachment of non-native species 
can all negatively impact plant and pollinator popula- 
tions, and may disrupt the interaction between them 
as well. For example, two recent reviews highlighted 
that increased fragmentation leads to an increased 
probability of pollinator failure and reduced plant 
reproductive success (Aizen et al. 2002; Wilcock and 
Neiland 2002). These effects can be exacerbated by 
increased inbreeding depression in small populations 
(e.g., Severns 2003; Kephart 2004). Even highly local 


245 


246 


fragmentation can reduce pollinator visitation rates and 
seed production, and lead to greater expression of in- 
breeding depression within metapopulations (Lennart- 
son 2002). 

Most of the aforementioned studies relating habitat 
loss to pollination issues focus on single plant species. 
It is essential, however, to first determine the relation- 
ships between interacting communities of plants and 
pollinators, so as to direct future research towards 
those interactions likely to be most important either 
for single-species conservation or for community-level 
restoration. An improved understanding of the repro- 
ductive ecology of native GOE forbs could lead to 
more effective management tactics. Thus, we investi- 
gated visitation patterns of potential insect pollinators 
to a forb community in a remnant of the GOE. Our 
goal was to determine the richness and abundance of 
visiting insect taxa and patterns of insect visitation to 
GOE forbs, in order to provide baseline data on plant- 
pollinator interactions within the GOE. Specifically 
we asked: 

(1) What forbs are present and what insects visit 
them? 

(2) How diverse are the insect visitors and do insect 
taxa visit only one plant species or many? 

(3) How do forb abundance and insect visitation 
change throughout the season? 


Methods 
Field Site 

The Cowichan Garry Oak Preserve near Duncan, 
British Columbia (48°48'30.8"N, 123°37'52.5"W) is 
a 12.3 ha remnant of the GOE which once encompassed 
as much as 45 000 ha throughout the Cowichan Val- 
ley. Within the private preserve (owned by the Nature 
Conservancy of Canada) there are a number of large, 
forb-dominated gaps in the woodland, one of which we 
chose for study. The Nature Conservancy of Canada 
actively manages the area, primarily through removal 
of C. scoparius. 


Forb surveys 

Prior to flowering in 2002, we laid out a 25 x 25m 
plot which encompassed the majority of the woodland 
gap. Within each 5 x 5 m cell of the plot, a 0.5 m? 
rectangular sampling quadrat was randomly located, 
for a total of 25 quadrats. The number of flowering 
stems was counted for each forb within each quadrat 
in six, weekly surveys from 23 April until 30 May. This 
period encompassed the majority of the flowering sea- 
son (see Results). Voucher specimens of all forbs were 
deposited in the Simon Fraser University Herbarium, 
Burnaby, British Columbia. Plant identification follows 
Douglas et al. (1998). For each of the forbs present, we 
also present estimates of flower size and flower number 
per inflorescence based on Douglas et al. (1998; Table 
1) as floral traits are known to affect insect visitation 
patterns. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Floral visitor surveys 

Several different methods can be used to assess the 
community of floral visitors, each with positive and 
negative attributes. Because we were interested in the 
interaction between plants and potential pollinators 
for the entire plant community, and wanted to impact 
the site as little as possible, we used a combination of 
direct observation and pan trapping for this research. 
Direct observation of visitors to flowers allows the best 
description of relationships between plants and poten- 
tial pollinators, and so was our method of choice. 
Direct observation of visitation, however, does not usu- 
ally allow complete identification of floral visitors “‘on 
the wing”. Survey methods like pan trapping or sweep 
netting can increase the total insect “catch” and allow 
for accurate identification of captured specimens, but 
information gained has limited utility for community- 
level studies as these methods do not allow evaluation 
of interactions between plants and visitors and fre- 
quently capture insects that are not floral visitors. In 
addition, sweep netting can damage plants by breaking 
off flowering stems. Because we wanted to increase 
our number of captured insects for identification, we 
used pan traps as the less damaging capture method. 
Floral visitor surveys were performed from 23 April 
to 23 May, and then were terminated as there were 
few flowering stems remaining. 


Direct observations 

Each of the 25 quadrats was observed for two, 15- 
minute periods each week by two observers. Observa- 
tions were made on sunny days between 09:00 and 
17:00, and the order of quadrat observation was ran- 
domised for each of the two observers. A visit was 
counted if an insect was observed probing at least one 
flower on a plant. Insect taxa were identified based on 
morphological features visible in the field (morphos- 
pecies) and representative specimens of each morphos- 
pecies were collected between observation periods and 
identified as completely as possible in the lab. Because 
of low visitation rates and our desire to minimize our 
impact on pollinator populations, we collected only 
enough specimens to confirm morphospecies. In most 
cases morphospecies contain several taxonomic species, 
a limitation of the direct observation method, and so 
our identifications to genus will under-represent the 
taxonomic diversity of floral visitors. For example, 
Osmia sp. | are large-bodied Osmia (approximately 
| cm long) and include O. lignaria, O. texana, and 
possibly other species with similar appearance on the 
wing, but not captured for inclusion with our voucher 
specimens. Because visitation rates were generally low 
(see Results), visits to flowering stems outside the 
quadrats were also recorded. These casual observations 
may have been biased towards larger, more apparent 
insects, but are included in an effort to make visitation 
data as complete as possible. 


2005 PARACHNOWITSCH AND ELLE: INSECT VISITATION 247 


Camassia quamash 

200 Cerastium semidecandrum* 
Collinsia parviflora 
Dodecatheon hendersonii 


i¢2) ; 
S Erythronium oregonum 
D 150 Lomatium utriculatum 
zs Ranunculus occidentalis 
i Vicia sativa * 
® 
= 100 
is) 
© 
Q 
& 50 
5 
Fas 

0 

110 120 130 140 150 


Mmm Andrena 
INN Apis * 
Bombus 

| _] Diptera 

{7 4A Lasioglossum 
HI] Osmia 


Number of visits within quadrats 


110 120 130 140 150 


Julian Date 


FicurE 1: A. (top) Phenology and abundance of visited plant species (within quadrats) at the Cowichan Garry Oak Preserve. 
B. (bottom) Phenology of insect visitation within quadrats grouped by genus of visitors (family for Diptera). An 
asterisk (*) indicates non-native species origin. No observations of floral visitors were made during the last floral 
survey due to the large number of quadrats with few or no flowering stems. 


248 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |. List of flowering plant species sampled in quadrats, followed by six other species that we observed flowering at 
the Cowichan Garry Oak Preserve but which were not present in quadrats. An asterisk (*) indicates non-native origin, and a 
dagger (7) indicates an endangered or threatened species. Peak flowering date includes the total number of flowering stems 
in all quadrats. Small, medium and large flower size estimates indicate flower diameters <lcm, 1-3 cm, and >3 cm, respec- 
tively. Flower number indicates plants that usually have few open flowers per stem (<10) or many (>10), unless indicated 


more precisely (Douglas et al. 1998). 


Scientific name 


Cardamine sp. 
Camassia quamash 
Cerastium arvense 


Cerastium semidecandrum* 


Claytonia perfoliata 
Collinsia parviflora 
Cytisus scoparius* 
Dodecatheon hendersonii 
Erythronium oregonum 
Galium aparine 
Geranium molle* 
Lathyrus sphaericus* 
Lithophragma parviflorum 
Lomatium utriculatum 
Montia linearis 

Myosotis discolor* 

Phlox gracilis 
Ranunculus occidentalis 
Sanicula crassicaulis 
Valerianella locusta* 
Vicia sativa* 


Also flowering at the site: 
Achillea millefolium Yarrow 
Fritillaria affinis 
Sericocarpus rigidus* 
Triteleia grandiflora var. howelliit Howell’s Brodiaea 
Viola praemorsa ssp. praemorsat 
Zygadenus venenosus 


Pan trapping 

A set of three pan traps (yellow, white, and blue to 
reduce colour bias: Leong and Thorp 1999) was placed 
at a random point along each of the five north-south 
transects making up the 25 x 25 m” grid (i.e., five sets). 
Pans were filled with ca. 250 mL of water and a few 
drops of glycerol (Kearns and Inouye 1993) and set out 
on each observation date for at least 6 hours. Cap- 
tured insects were preserved in ethanol prior to iden- 
tification. Identification of Hymenoptera, Lepidoptera 
and Diptera followed Michener et al. (1994), Guppy 
and Shepard (2001) and McAlpine et al. (1981), res- 
pectively. Voucher specimens of all morphospecies 
were deposited with the Simon Fraser University 
Natural History Museum. 


Statistical analyses 

To illustrate the overall patterns of visitation, we 
present a matrix of the interactions between forbs and 
putative pollinator morphospecies (Table 1) and cal- 
culate the sum of interaction types for both the forb 
species and insect morphospecies. For each sampling 


Peak flowering Flower Flower 
Common name date (# stems) size number 
Bitter-cress 23/04/02 (12) small few 
Common Camas 16/05/02 (112) large 5+ 
Field Chickweed 31/05/02 (6) medium few 
~ Little Mouse-ear 23/04/02 (25) small few 

Miner’s-lettuce 01/05/02 (13) small few 
Blue-eyed Mary 01/05/02 (196) small | — many 
Scotch Broom 23/05/02 (3) medium many 
Broad-leaved Shootingstar 23/04/02 (62) medium 2-15 
White Fawn Lily 23/04/02 (7) large usually | 
Cleavers 16/05/02 (600) small many 
Dovefoot Geranium 23/05/02 (15) medium 2 — many 
Grass Peavine 23/05/02 (37) medium few 
Small-flowered Woodland Star 01/05/02 (1) medium 5-11 
Spring Gold 16/05/02 (63) small many 
Narrow-leaved Montia 23/04/02 (4) small many 
Common Forget-me-not 31/05/02 (4) small many 
Slender Phlox 16/05/02 (14) small many 
Western Buttercup 08/05/02 (56) medium few 
Pacific Sanicle 23/05/02 (3) small many 
Corn Salad 16/05/02 (758) small many 
Common Vetch 31/05/02 (63) medium few 

small many 
Chocolate Lily large 1-5 
White-top Aster small many 

medium many 
Yellow Montane Violet medium few 
Meadow Death-camas medium many 


date, we used a x? analysis to test whether the number — 
of visits to each plant species (summed across quadrats) — 
was proportional to the number of available stems of — 
each species. All plant species that were flowering | 
within quadrats on a particular date were included in 
these analyses. To test whether visitation patterns were | 
determined by the phenology of the forbs (i.e., that | 
later-flowering species had more visits), we performed | 
a regression of the total number of visits a plant species . 
received (pooled over all dates and quadrats) on peak | 
species flowering date (1) including all species, and | 
(2) including only the eight forbs that received visits, 
to avoid bias due to the many zero values for unvisited 
species. 


Results 

Twenty-one forb species were observed flowering 
within quadrats, 14 native and 7 non-native (Table 1). 
Most of the species had small (11 species) or medium 
(8 species) flower size while only Camassia quamash 
and Erythronium oregonum had large flowers (greater | 
than 3 cm diameter); 10 of the species generally pro- 


249 


INSECT VISITATION 


PARACHNOWITSCH AND ELLE 


2005 


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duce fewer than 10 flowers per flowering stem (Table 
1). Only 10 of the forb species had a density greater 
than 2 flowering stems/m? in our quadrats on their peak 
flowering date (Table 1). Our sampling period cap- 
tured the peak spring flowering period of 13 of these 
species (Table 1, Figure 1A); five species had a peak 
flowering date before the start of the survey (or on 23 
April), including Cardamine sp., Cerastium semidecan- 
drum, Dodecatheon hendersonii, Erythronium oregon- 
um and Montia linearis while only Cerastium arvense, 
Myosotis discolor, and Vicia sativa continued to in- 
crease in abundance after the survey. Six additional 
native forb species were present at the site, including 
three species at risk, but were not present in our sam- 
pling quadrats due to low frequency or patchy distri- 
bution (Table 1). 

Visitation frequency was low (0.5-3.9 visits/hr). 
Flower visitors were observed on 11 forb species 
(Table 2) but on only 8 of the 21 forbs available with- 
in the quadrats (Table 3). On four of the five sampling 
dates the number of visits to each forb species was 
different from that expected based on the number of 
flowering stems available (23 April : x =28.7, df = 8, 
P =0.0004; 1 May: x? = 12.8, df = 13, P=0.4666; 8 May: 
° =01017,.di= 11? < 0.0001) te May-47 =312.7, 
df= 15; P< 0/0001; 23 May? x? = 182 1 dt=16, P< 
0.0001). 

Thirteen insect morphospecies were observed vis- 
iting forbs flowering within quadrats, and eight addi- 
tional morphospecies were observed visiting forbs at 
the site (Table 4). Hymenoptera accounted for the 
largest percentage of visits within quadrats (71.4%) 
with the remaining visits by Diptera. Osmia sp. 1 were 
large-bodied mason bees, Osmia sp. 2 were small-bod- 
ied mason bees; Andrena morphospecies were distin- 
guished based on relative size, abdominal markings, 
and dense hairs on the thorax. “Other Syrphidae” refers 
to the common, striped hoverflies such as species of 
Syrphus and Metasyrphus which could not be distin- 
guished on the wing. The majority of observed visits 
were made by Apis mellifera (honeybee), followed by 
the native Osmia and Andrena spp. (Table 4). Within 
quadrats, A. mellifera was observed visiting C. qua- 
mash exclusively, and 88.9% of observed A. mellifera 
visits occurred during the date of peak C. quamash 
bloom. Most floral visitors did not exhibit this fidelity, 
however; 13 of the 21 morphospecies observed visited 
more than one forb species, and eight of the 11 visited 
forb species were visited by more than one insect mor- 
phospecies (Table 2). As the season progressed, more 
visits from a greater number of insect morphospecies 
were observed until most of the forbs had senesced 
(Figure 1B). Although there were more visits observed 
later in the season, there was no relationship between 
peak flowering date and the number of visits received 
by a plant species (R* = 0.0002, F, ;, = 0.00, P = 0.96), 
even when only the eight visited forbs were consid- 
etedi(h’ = 0.079; FO 375 0 30): 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE 3. Number of floral visits observed within quadrats 
by insect morphospecies. Visits are summed across 
quadrats, the eight visited plant species and surveys from 
23 April to 23 May 2002. 


Floral visitors # Visits % of 
Visits 

Hymenoptera 
Andrena sp. | ] 1.4 
Andrena sp. 2 9 Wd.) 
Andrena sp. 3 4 527 
Apis mellifera 18 25g 
Bombus bifarius 4 Sif 
Bombus flavifrons ] 1.4 
Lassioglossum sp. z p25) 
Osmia sp. | 10 14.3 
Osmia sp. (male) 1 1.4 

Diptera 
Bombylius major 6 8.6 
Unidentified bombyliid sp. 3 4.3 
Volucella bombylans 2 29 
Other Syrphidae 2) 22 


TABLE 4. Distribution of observed visits (all insect species 
combined) among the 8 plant species that received visits 
within quadrats. Data derived by pooling across all quadrats 
and observation periods (23 April to 23 May, 2002). An 
asterisk (*) indicates non-native origin. 
# Visits % of 
Visits 


Plant species 


Camassia quamash 37 
Cerastium semidecandrum* l 
Collinsia parviflora 8 
Dodecatheon hendersonii 5 71 
Erythronium oregonum | 
Lomatium utriculatum 7 
Ranunculus occidentalis 10 
Vicia sativa* 


Pan traps caught only nine individuals throughout 
the sampling period, including four Andrena (species 
2, 3 and one additional morphospecies not seen dur- 
ing the observation periods), one Osmia male (a dif- 
ferent morphospecies than the male Osmia seen dur- 
ing the observation period), one Volucella bombylans, 
two of the unidentified Bombyliid observed visiting 
flowers, and a male Megachile sp. that was not ob- 
served visiting flowers. Low catches precluded further 
analysis. 


Discussion 

Visitation rates within this fragment of the GOE 
were low in 2002 (maximum rate 3.9 visits/hr of obser- 
vation, on 16 May), but eight of 11 forb species that 
received visits were visited by more than one insect 
morphospecies and 13 of the 21 insect morphospecies 
were observed visiting more than one plant species 
(Table 2). Because of the low number of visitors ob- 
served, it is difficult to make inferences about the 


2005 


degree of specialisation of individual insect morphos- 
pecies; however, most visited forbs appear to have 
generalised insect visitation patterns. Similarly gen- 
eralised visitation patterns were observed at the same 
site in 2001, although visitation rates were higher (Elle 
and Carney 2003). In that survey, conducted over a 
shorter time (equivalent to our third and fourth census 
dates) and so not inclusive of the flowering period of 
many GOE forbs, visitation by 14 insect morphospecies 
to just four forb species (all visited by multiple mor- 
phospecies) was observed at a rate of 28.5 visits/hr. 
Among-year variation in visitation rate is not uncom- 
mon in spring-flowering communities (e.g., Motten 
1986; Burd 1994), but it is unknown whether the un- 
predictability of visitation limits reproductive suc- 
cess of GOE forbs. 

The number of observed visits to each forb species 
differed from that expected based on flowering stem 
availability on all but one sampling date, suggesting 
some degree of preference on the part of insect visitors. 
Camassia quamash was visited by the most diverse 
group of insects (16 of the 21 observed morphospecies) 
and received the majority of visits (52.9 %). On its 
peak flowering date (16 May), C. quamash was com- 
mon (flowering in 22 of 25 quadrats), and received 
24 of the 26 observed visits. This forb produces large 
flowers on multi-flowered inflorescences (Table 1), and 
may have high nectar rewards. In contrast, the majority 
of forbs within our quadrats with no observed visits (9 
of 11) have small and/or few flowers (Table 1), and may 
be less attractive to insect foragers; several are likely 
self-fertilised (1.e., Galium aparine, Montia linearis, 
Myosotis discolor). Although low visitation rates tem- 
per our conclusion of non-random visitation, similarly 
non-random visitation was observed at this site the year 
before the current study (Elle and Carney 2003). 

The composition of the flowering forb community 
and the insects that visited them changed over the dura- 
tion of our study (Figure 1). It is possible that the phe- 
nological differences in forb abundance influenced 
visitation. For instance, C. quamash was the most 
abundant plant on 16 May, which coincided with the 
greatest number of observed visits, in particular by Apis 
mellifera, which visited C. quamash almost exclusively 
(Figure 1B). However, the results of the regressions of 
number of insect visits on peak flowering date sug- 
gest that flowering later does not lead to increased total 
visitation; we suggest instead that C. guamash is sim- 
ply highly attractive to insects. 

The high abundance and diversity of visitors to C. 
quamash (at least relative to other plants) suggests 
that it may have an important ecological role in this 
community; it may serve as a “magnet” species for 
pollinators and enhance pollination of neighbouring 
species by increasing the overall size and/or diversity 
of the pollinator community (Thomson 1978; Laverty 
1992). The only pollinator species that were present 
but not observed visiting C. guamash included rarely 
observed species (two Lepidoptera; Table 2) and pos- 


PARACHNOWITSCH AND ELLE: INSECT VISITATION 


251 


sibly a specialist fly (the unidentified Bombyliid was 
observed numerous times outside the quadrats but only 
visiting Lomatium utriculatum). Our study did not ad- 
dress interactions of C. guamash with other plants in 
the community; however, the relatively high frequency 
and variety of visitors to C. guamash suggests that future 
research could test whether this species has competitive 
or facilitative effects on forb pollination within the GOE. 

Bees were the most common floral visitors in our 
study and are major pollinators globally (Kearns and 
Inouye 1997). Early in the sampling period, Osmia and 
Andrena spp. were frequently observed (Figure 1B), 
although the most frequently observed visitor overall 
was Apis mellifera (honeybee), which has been intro- 
duced world wide from Europe and is used for crop 
pollination (Kearns and Inouye 1997). Apis mellifera 
has been implicated in the decline of native pollinators 
(Kearns et al. 1998 and references therein; Goulson 
2003; Thomson 2004) and so its potential impacts on 
plant-pollinator relationships in the GOE need to be 
considered. Our study did not address interactions be- 
tween A. mellifera and other bees. However, A. mellif- 
era visited C. quamash almost exclusively both in the 
present study and a previous study at this site (Elle 
and Carney 2003), and thus the potential for negative 
effects on other pollinators will depend on the impor- 
tance of C. guamash as a nectar or pollen resource for 
native pollinators (which generally visited other plants 
in addition to C. quamash, Table 2). Our data do sug- 
gest that if native pollinators were to decline for rea- 
sons other than competition with honeybees (e.g., loss 
of appropriate nesting habitat), A. mellifera would not 
provide pollination services for plant species other 
than C. quamash. 

Our study provides initial data on plant-pollinator 
interactions in the GOE, but additional, detailed study 
is necessary to determine the importance of plant- 
pollinator mutualisms for the sustainability of the com- 
munity as a whole. Further research is needed to deter- 
mine the effectiveness and importance of insect visitors 
for pollination of GOE forbs, and whether these insects 
are at risk due to habitat or nesting requirements. The 
numerous forb species for which we’ve never observed 
any visits, at both the Cowichan Garry Oak Preserve 
and other sites (Elle and Carney 2003) should be inves- 
tigated to determine if they are, indeed, reproducing 
maximally via autogamy, or if they are instead pollen 
limited. Our data suggest that pollination of visited 
plant species may be generalized, however, and so this 
ecosystem may be resilient to the loss of individual 
pollinating insect species (Waser et al. 1996). More 
research is important to determine whether rare GOE 
forbs are at risk due to failed mutualisms (e.g., Bond 
1994). 


Acknowledgments 

The authors thank T. Ennis and the Nature Conser- 
vancy of Canada for access to the Cowichan Garry Oak 
Preserve. J. Biernaskie, S. Campbell, C. Caruso, B. 


252 


Husband, M. Schuetz and two anonymous reviewers 
provided constructive feedback on earlier versions of 
the manuscript, and J. Biernaskie, M. Schuetz, and M. 
Thane provided field assistance. Funding was provided 
through an Undergraduate Student Research Award 
to A.L.P. and an operating grant to E.E., both from the 
Natural Sciences and Engineering Research Council 
of Canada. 


Documents Cited (marked * in text) 
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goert.ca. Accessed online May 2004. 


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Received 18 October 2002 
Accepted 5 April 2005 


An Annotated Checklist of the Spiders of Newfoundland 


J. R. PICKAVANCE! and C. D. DONDALE? 


'Biology Department, Memorial University, St. John’s, Newfoundland and Labrador A1B 3X9 Canada; rpickava@mun.ca. 
Eastern Cereal and Oilseed Research Centre, K. W. Neatby Building, Central Experimental Farm, Ottawa Ontario K1A 0C6 


Canada 


Pickavance, J. R., and C. D. Dondale. 2005. An annotated checklist of the spiders of Newfoundland. Canadian Field-Natu- 


ralist 119(2): 254-275. 


Since Hackman’s report in 1954 of 216 (adjusted from a total of 220 for apparent errors) spider species from Newfound- 
land, the total has grown by 68% to 363. An annotated checklist is presented. Of this total, 223 (c. 61%) are Nearctic, 119 
(c. 33%) are Holarctic and 21 (c. 6%) are introduced. Comparisons are made with other Canadian spider faunas. The intro- 
duced species and the proportions of Linyphiidae, Lycosidae and Dictynidae are discussed. 


Key Words: Araneae, species distribution, introduced species, Newfoundland. 


Hackman’s (1954) list of Newfoundland (referring 
solely to the insular portion of the province of that 
name) spiders was the first such report for any Canadi- 
an province. Since then lists for four more provinces 
and territories have been published: British Columbia 
(West et al. 1984, 1988; Bennett 2001); Yukon (Dondale 
et al. 1997); Manitoba (Aitchison-Benell and Dondale 
1990), and Quebec (Bélanger and Hutchinson 1992; 
Paquin et al. 2001, Paquin and Dupérré 2003). We have 
made use of additional unpublished data for Alberta and 
Saskatchewan (Buckle and Holmberg 2004*) and 
British Columbia (Bennett et al. 2004*). 

In 2002 the formal, political name of the province 
formerly called Newfoundland was changed by Act of 
Federal Parliament to Newfoundland and Labrador. 
Therefore since that date it has been proper to use 
Newfoundland to refer solely to the insular part of the 
province, and that is the practice adopted here. New- 
foundland was entirely or almost entirely covered by 
ice in the most recent (Wisconsin) glaciation, which 
reached its maximum extent probably around 24 000 
years BP (Dyke et al. 2002). The ice then underwent 
a series of retreats until the coastal margins of New- 
foundland were exposed around 12 000 BP (Shaw 
2003), after which time the ice continued to retreat and 
leave the land exposed. The modern fauna was presum- 
ably established by post-glacial immigration aided by 
some anthropogenic introductions. The presence or ab- 
sence of biological refugia during that glaciation has 
long been debated. Such refugia could have been im- 
portant contributors to the post-glacial colonisation, 
and evidence thought to indicate their existence in 
Newfoundland has been presented from a variety of 
taxa (e.g., Fernald 1924; Wynne-Edwards 1937; Bel- 
land 1987; Hamilton and Langor 1987). Ballooning 
from continental North America on the region’s prevail- 
ing westerly winds probably accounted for the post- 
glacial arrival of most of Newfoundland’s spider species. 


It is not altogether clear which species were brought 
to Newfoundland by humans. The introduction of Palae- 
arctic species by humans is relatively straightforward. 
The first documented European visitors were Vikings, 
briefly established at L’Anse aux Meadows around 
1000 BP (Wallace 1991). Then since the late 15" cen- 
tury European settlement and commercial traffic can 
explain the presence of Palaearctic species in New- 
foundland. In addition, some Nearctic and Holarctic 
species may have been brought to the island by the traf- 
fic between Newfoundland and the maritime provinces, 
Labrador and the eastern seaboard of the United States. 
There were also approximately 5 000 years of aborig- 
inal occupation of Newfoundland before the arrival of 
Europeans, with continual aboriginal traffic between 
Labrador and Newfoundland (Renouf 1999). This may 
account for the presence of some Holarctic and sub- 
arctic-Nearctic species. 

By whatever means of arrival, 363 species are now 
known from Newfoundland, an increase of 68% over 
Hackman’s (1954) report of 216 (adjusted total) spe- 
cies. 

Ecology of Newfoundland 

Newfoundland lies in the northwest Atlantic between 
approximately 47°-52° north and 53°-59° west and lies 
within the general ecological region of the Boreal 
Shield Ecozone. Eleven Ecoregions (encompassing 
25 Ecodistricts) are distinguished in Newfoundland, 
and full details of these are in Ecological Stratification 
Working Group (1995). More general biogeographical 
information can be found in South (1983). 

Newfoundland’s climate is influenced by the sur- 
rounding Atlantic ocean in general and the cold Labra- 
dor current to the east in particular. Climate varies 
between the Ecoregions, from a low subarctic ecocli- 
mate in the Strait of Belle Isle Ecoregion on the North- 
erm Peninsula (mean annual temperature 2.5°C), through 
the maritime high boreal ecoclimate of the Long Range 


254 


2005 


Mountains (mean annual temperature 4°C), the mar- 
itime mid-boreal ecoclimate of Central Newfoundland 
(mean annual temperature 4.5°C), to the oceanic low 
boreal ecoclimate of the South Avalon-Burin Oceanic 
Barrens (mean annual temperature 5.5°C). The domi- 
nant vegetation is a mixture of Black Spruce (Picea 
mariana), White Spruce (Picea glauca), Balsam Fir 
(Abies balsamia), Tamarack (Larix decidua), lesser 
amounts of birch (Betula), extensive peatlands, com- 
munities of ericaceous shrubs, crowberry (Empetrum) 
barrens, and mosses and lichens. At higher elevations 
areas of semi-exposed bedrock are frequent. 


Materials and Methods 
Abbreviations 

Throughout the text the following abbreviations may 
be employed: AB = Alberta; AMNH = American Mus- 
eum of Natural History; BC = British Columbia; BL 
= Belcher Islands, Hudson Bay; CDD = C. D. Dondale; 
CNC = Canadian National Collection of Insects and 
Arachnids, Agriculture and Agri-Food Canada, Ottawa; 
GL = Greenland; GMNP = Gros Morne National Park, 
Newfoundland; FMNH = Finnish Museum of Natural 
History, Helsinki; HZ = Hazen Camp, Ellesmere Is- 
land; JRP = J. R. Pickavance; MB = Manitoba; MCZ 
= Museum of Comparative Zoology, Harvard; NF = 
Island of Newfoundland; PL = Peary Land, Greenland; 
QC = Quebec; SK = Saskatchewan; TNNP = Terra 
Nova National Park, NF; YK = Yukon. 


A note on Hackman’s species total 

We have reduced Hackman’s (1954) total of 220 by 
four to give a revised total of 216. Hackman (1954) 
listed some species on the basis of immature specimens, 
_ some of which have not subsequently been confirmed. 
~CDD examined the supposed juveniles of Tetragnatha 
caudata Emerton and T: vermiformis Emerton and con- 
cluded that the specimens are of uncertain identity. 
These two species have therefore been removed from 
Hackman’s (1954) total. Hackman (1954) recorded 
three species of Dolomedes from Newfoundland: D. 
fulviatronotatus Bishop, D. scapularis C. L. Koch and 
D. vittatus Walckenaer. Dolomedes fulviatronotatus is 
now D. striatus Giebel, and his D. vittatus was a mis- 
identified specimen of D. striatus. In addition, his 
record of D. scapularis (junior synonym of D. triton 
(Walckenaer)) cannot be confirmed because this species 
is not in Hackman’s collection in FMNH, has not 
subsequently been found in Newfoundland, and has 
never been brought in by the public for identification 
(unlike virtually all other species with large adults). 
We have therefore removed two of the three Dolo- 
medes from Hackman’s (1954) total. 

This paper is based on published reports of New- 
foundland spiders, specimens in the CNC, examination 
by CDD of collections of Newfoundland spiders de- 
posited elsewhere, and extensive collections by JRP 
over the last 14 years. Apart from the major collections 

reported by Hackman (1954), reports of Newfoundland 


PICKAVANCE AND DONDALE: SPIDERS of NEWFOUNDLAND 


255 


spiders are rare. These consist of accounts of scattered, 
small collections (e.g., Pickard-Cambridge 1881; Emer- 
ton 1914, 1915, 1927) or lists assembled for particular 
purposes, e.g., Lindroth’s (1957) comparison of Euro- 
pean and North American faunas. A number of col- 
lectors (e.g., Lloyd Hollett and Kevin Pardy) have 
deposited Newfoundland specimens in the CNC but 
have not formally reported on their collections. Most 
records of Newfoundland spiders occur in taxonomic 
works such as Gertsch and Ivie (1955), Ivie (1969), 
Leech (1972), Dondale and Redner (1978, 1982, 1990), 
Platnick and Dondale (1992), Buckle and Roney (1995), 
Saaristo and Koponen (1998), and Miller (1999). 

We have followed Platnick (2005) for familial place- 
ment for the sake of uniformity and consistency. We 
have followed Platnick (2005) for genera and species 
names, except that we follow Buckle et al. (2001) for 
linyphiid nomenclature. For convenience families 
and species are in alphabetical order. 

Species recorded by Hackman (1954) are indicat- 
ed “[H]’. Included in such brackets is the name he 
used if different from the name in Platnick (2005*) 
and other relevant notes. 

The designations of Nearctic, Holarctic, or Palae- 
arctic listed for each species have been gleaned from 
various sources (e.g., Buckle et al. 2001; Dondale and 
Redner 1978, 1982, 1990; Dondale et al. 2003; Plat- 
nick 2005*; Roberts 1993). Here we use these terms to 
mean the biogeographical origin of a species rather 
than the present-day distribution. For example, Araneus 
diadematus Clerck is listed as Holarctic by Platnick 
(2005*), but is originally a Palaearctic species intro- 
duced to North America (Dondale et al. 2003). There- 
fore we refer to this and similar species as ““Palaearctic; 
introduced”. 

It has sometimes been difficult to decide whether a 
species should be called Palaearctic introduced or truly 
Holarctic. For example, Buckle et al. (2001) describe 
Erigone dentipalpis (Wider in Reuss) as “Introduced?” 
while Platnick (2005*) lists it as Holarctic. Pending 
clarification we treat this and similar cases as intro- 
duced Palaearctic species. Spider introductions to North 
America were also dealt with by Lindroth (1957), but 
because significant portions of his information about 
spiders has been superceded by later work we have 
placed less emphasis on his records. A particularly 
problematic species is Theridiosoma gemmosum (L. 
Koch), traditionally regarded as Palaearctic introduced 
to North America (e.g., Locket and Millidge 1953). 
Coddington (1986) questioned this and pointed out it 
might be a Nearctic species introduced to Europe be- 
cause it is common in North America but rare in Eu- 
rope. Here we follow the traditional position because 
the species is confined to the eastern part of North 
America, where it may be common because of the 
amount of its preferred habitat of old-growth and (to a 
lesser extent) second-growth forest (Coddington 1986), 
whereas it is very widespread in Europe and Asia. 


256 


The true distribution and habitat preferences of most 
Newfoundland spider species are unknown, with some 
species known only from one or two records. Note 
that the localities in the list below more often reflect 
the activities of collectors than the true distribution 
of the species. Only a general habitat description is 
given for each species unless there is some obvious 
correlation with a particular environment. Note that the 
term “mixed coniferous woods” (which occurs regu- 
larly in the checklist) encompasses a wide diversity 
of micro-habitats such as dry litter, wet litter, patches 
of wetland, moss, lichens, bark, leaves, standing water, 
running water, patches of hardwood shrubs and patch- 
es of herbs as well as boundaries with more extensive 
areas of wetland, barrens or rocky areas. Additional 
habitat information has been taken from Bélanger 
and Hutchinson (1992) because the majority of New- 
foundland species occur in Quebec, which at least in 
the middle of that province is broadly ecologically 
similar to Newfoundland. Habitat information has also 
been gathered from a variety of other sources, e.g., 
Chamberlin and Gertsch (1958); Dondale and Redner 
(1978, 1982, 1990); Dondale et al. (2003); Levi (1971); 
Opell and Beatty (1976); Paquin and Dupérré (2003) 
and Platnick and Dondale (1992). 


Dates, Depositories and Localities 

Dates are expressed e.g. 5Aug98; an unknown day 
and/or month indicated by 00 (but 00 in the year posi- 
tion means 2000); specimens from a range of dates are 
indicated e.g. 12-28Aug98. If no date is given collec- 
tion date is uncertain. One specimen is indicated Go 
or 9; more than one specimen COC or 9 9. Records 
from other collectors are not included except where no 
records exist in the JRP collection. All specimens are 
in the JRP collection (Biology Department, Memorial 
University of Newfoundland) unless otherwise noted; 
specimens held elsewhere are indicated e.g. (CNC); 
specimens of species held both in JRP collection and 
elsewhere are indicated e.g. (and CNC). For each spe- 
cies a maximum of five locations is given; if a species 
is known from more than five locations, five were 
selected to show the general distribution. Locations 
are arranged alphabetically. Place names are in accor- 
dance with Natural Resources Canada: Canadian Geo- 
graphical Names (2005*) where additional data such 
as latitude and longitude can be found. 


Checklist of Newfoundland Spiders 
AGELENIDAE (4 spp.) 

Agelenopsis utahana (Chamberlin & Ivie, 1933) [H] 
Nearctic. Among herbaceous plants in clearings in coniferous 
woods; corners of walls and fences. Norris Point 18Aug 029°; 
Port au Choix 12Jul99¢C. 


Tegenaria atrica C. L. Koch, 1843 

Palaearctic; introduced. Warehouses in Donovans Industrial 
Park, St. John’s. 

St. John’s OOMay95°. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Tegenaria domestica (Clerck, 1758) 

Palaearctic; introduced. This synanthropic species is common 
across Newfoundland inside houses (particularly basements), 
sheds and other structures. 

Norris Pointl4Aug979; Port au Choix 9Jul04¢; Portugal 
Cove 27May89°9;; St. John’s 29Sep97° ; TNNP 1 1Jun93¢. 


Tegenaria duellica Simon, 1875 

Palaearctic; introduced. Buildings in the Waterford Bridge 
valley, St. John’s. 

St. John’s 1Jun0O1°; St. John’s 20Sep04c¢. 


AMAUROBIIDAE (5 spp.) 

Amaurobius borealis Emerton, 1909 [H; Walmus borealis 
(Emerton) | 

Nearctic. Litter of mixed coniferous woods. 

Berry Head Pond (GMNP) 10Jul00°; Gander 00Aug98¢; 
Glide Lake 00Jul96° ; Main River west 29Jul00¢. 


Callobius bennetti (Blackwall, 1846) [H] 

Nearctic. Litter of mixed coniferous woods; among stones 
at the back of beaches. 

Bakers Brook estuary 11Aug99°; Green Point 26Aug98° ; 
Port au Choix 5Jul04°; Sugar Loaf (St. John’s) O0OOct92°2 ; 
TNNP 11Jun93¢. 


Coras montanus (Emerton, 1890) [H; Agelenidae] 

Nearctic. Litter of mixed coniferous woods; under bark; in 
crevices between rocks. 

Harbour Grace 27May94° ; St. John’s OO0Jan96° ; St. John’s 
OOSep97¢. 


Cybaeopsis euopla (Bishop & Crosby, 1935) [H; Callioplus 
euoplus Bishop & Crosby] 

Nearctic. Litter of mixed coniferous woods. 

Berry Head Pond (GMNP) 19Jul00°; Glide Lake 00Jul9609 
; Main River west OO0Jul98°9; Nameless Cove 16Aug01 9° ° 
; Port au Choix 8Jul00¢. 


Cybaeopsis tibialis (Emerton, 1888) [H; Callioplus tibialis © 
(Emerton)] j 
Nearctic. Litter of mixed coniferous woods. 
Main River west 6Jun98 °c. 


ARANEIDAE (20 spp.) 

Aculepeira carbonarioides (Keyserling, 1892) 

Holarctic. In NF, known only from higher elevations of the 
Long Range Mountains. It spins large webs strung between 
boulders and is often seen at the centre of the web during 
daylight. | 
Gros Morne Mountain 14Aug94°; Highlands of St. John 
1Aug98° ; Killdevil Mountain 9Aug93° ; Tablelands 12Jun94° 
?; Winter House Brook canyon 31Jul97? °c. 


Araneus corticarius (Emerton, 1884) [H; from a juvenile | 
female that is not A. corticarius| . 
Nearctic. Mixed coniferous woods; among bog and swamp- | 
loving trees and shrubs. 

Main River west 12Aug00¢. 


Araneus diadematus Clerck, 1758 [H] 
Palaearctic; introduced. This is the common Garden Spider of | 
western Europe, often called the Cross Spider in North Amer- — 
ica because of the characteristic dorsum pattern. Widespread | 


in NF, particularly abundant around human settlements and |} 


associated gardens. Also found away from human habitations, | 
although this often indicates an abandoned community (see 
the discussion of this species below). 

Lewisporte 00Oct92° ; Mortier Bay O0Oct92° ; Norris Point 
18Aug999G; Paradise 0D0Aug93°; St. John’s 8Sep98°. 


2005 


Araneus groenlandicola (Strand, 1906) 

Nearctic. On herbs, low shrubs and stunted trees on open or 
barren land near the coast. In NF known only from coastal 
Northern Peninsula; it may be confined to that colder, sub- 
arctic area. 

Bakers Brook estuary 27Aug98°; Burnt Cape 4Aug98° ; 
Point Riche 15Aug97° ; Port au Choix 20Aug02° ° ; Shallow 
Bay 9Aug98° °. 


Araneus marmoreus Clerck, 1758 

Holarctic. Mixed coniferous woods; low shrubs near such 
woods; on rock-cuts along roads through such woods. 
Mount Scio 00Aug95¢; Norris Point 30 Jul97°; Rocky 
Harbour (Millbrook) 8Aug97° ; Woody Point (Lookout Hills) 
11Aug97°. 


Araneus nordmanni (Thorell, 1870) 

Holarctic. Mixed coniferous woods; wooden fences and break- 
waters. 

Boutitou 17Jul03°; Ferryland 21Sep98°. 


Araneus saevus (L. Koch, 1872) [H; Araneus solitarius 
(Emerton 1884)] 

Holarctic. Mixed coniferous woods; garden vegetation. 
Topsail (St. John’s) I5Aug96¢. 


Araneus trifolium (Hentz, 1847) [H] 

Nearctic. Low shrubs on open or barren ground; tall herbs in 
clearings in coniferous woods. 

Burgeo 13Aug03°; Lobster Cove 20Aug99°; Logy Bay 
20Aug03°; Norris Point 15Jul99¢; Rocky Harbour (Bot- 
tom Brook bog) 16Jul99¢. 


Araneus washingtoni Levi, 1971 

Nearctic. Mixed coniferous woods; low bushes. 

Eddies Cove West 30Jul49° (MCZ?); Gambo 25Apr49° 
(MCZ?). 


_ Araniella displicata (Hentz, 1847) [H; Araneus displicatus 
(Hentz) | 

Holarctic. Mixed coniferous woods. Common. 

Bakers Brook estuary 12Aug97°; Gander O0Jul98¢9; Logy 
Bay 13 Aug 01°; Millertown 26Jun80° 9; St. John’s 8Sep98° 
fo) 


Araniella proxima (Kulczynski, 1885) 

| Holarctic. Mixed coniferous woods. In NF known only from 
| the west of the island. 

} St Lunaire 15Aug00¢. 


| Cyclosa conica (Pallas, 1772) [H] 

| Holarctic. Mixed coniferous woods. Common. 

} Blackhead (St. John’s) 26Jun972; Boutitou 17Jul03° ; Gander 
| 20Jul9899; St. John’s 13Sep98° ° ; Stuckless Pond 20Jul93° °. 


| Hypsosinga pygmaea (Sundevall, 1831) [H:; Singa variabilis 
| Emerton 1884] 

} Holarctic. Edges of mixed coniferous woods; shrubs and herbs 
| in peatlands. 

|} Bay d’Espoir 11Jul85° (CNC); Bottom Brook (which one 
junknown) 19Aug86° (CNC); North Arm Mountain (Bonne 
| Bay) 13Aug81° ? (CNC). 


Hypsosinga rubens (Hentz, 1847) 

|Nearctic. Mixed coniferous woods; shrubs and herbs in 
| peatlands. 

Gander O0O0Jun-O00Jul81S (CNC); Hinds Point 29Jun872 


PICKAVANCE AND DONDALE: SPIDERS of NEWFOUNDLAND 


ZT 


Larinioides cornutus (Clerck, 1758) 

Holarctic. Shrubs; low trees; between boulders on the Long 
Range Mountains (where it can co-occur with Aculepeira 
carbonarioides). Common. 

Big Brook 18Jun02° °; New Ferolle 13Jul04° 9; Quidi Vidi 
19Sep98° °; Tablelands 31Jul97° °; Table Point 16Aug99° 
9 


Larinioides patagiatus (Clerck, 1758) 

Holarctic. Mixed coniferous woods; low cliffs; tall herbs and 
shrubs; stunted trees; houses, outbuildings; lower elevations 
of mountain slopes. Common at least in the centre and west. 
Badger 24Jun809; Big Brook 25Jun02°; Boutitou 17Jul039; 
Norris Point 30 Jul97° ; Point Riche 15Aug97°2¢. 


Larinioides sclopetarius (Clerck, 1758) 

Palaearctic; introduced. On houses, sheds and outbuildings, 
fences, bridges; on mixed vegetation near such structures. 
Common. 

Deer Lake 12Sep98 °; Epworth 2Jun01°; Norris Point 
5Aug99°9; Rocky Harbour 9Aug97° ; St. John’s 4Nov93¢. 


Metepeira palustris Chamberlin & Ivie, 1942 [H] 

Nearctic. Clearings and fallen timber in mixed coniferous 
woods. 

Gander 28Jul98¢. 


Zygiella atrica (C. L. Koch, 1845) 

Palaearctic; introduced. Among rocks near the coast; on break- 
waters and other coastal structures; fences and buildings. 
Common around the greater St. John’s area and the Avalon 
Peninsula in general; also in Deer lake. Elsewhere around 
the coast only Z. nearctica is found. 

Deer Lake 12Sep97° ° ; Ferryland 21Sep98°°; Flatrock 2Nov 
97°39; Logy Bay 16Sep03°°; Quidi Vidi 12Oct97° °c. 


Zygiella nearctica Gertsch, 1964 [H; Zygiella montana (C. 
L. Koch, 1834)] 

Nearctic. Mixed coniferous woods; shrubs; low sea-cliffs; 
boulders, wharves and other structures near the ocean. 
Conche 16Jul03°°; Lomond 18Jul97¢; Port au Choix 
1Aug97° 2S; Table Point 5SAug97°.9; TNNP 1 1Jun93¢. 


CLUBIONIDAE (14 spp.) 

Clubiona abbotii L. Koch, 1866 [H] 

Nearctic. Litter of mixed coniferous woods; litter under shrubs; 
edges of peatlands. 

Bakers Brook estuary 8-27Aug00¢. 


Clubiona bryantae Gertsch, 1941 [H] 

Nearctic. Litter of mixed coniferous woods and shrubs; rocks 
at back of beaches; litter at edge of sand dunes; bogs. 

Berry Head Pond (GMNP) 19Jul00°; Blackhead (St. John’s) 
12Nov02°°°; Green Point 6Jul97¢; Nameless Cove 
16Aug01° °; Shallow Bay 14Aug98°. 


Clubiona canadensis Emerton, 1890 [H] 

Nearctic. Mixed coniferous woods; among herbs in clear- 
ings in woods; under bark; on sedges and herbs in wetlands; 
in leaf-litter in gardens and cultivated areas. Common. 
Bakers Brook 8-27Aug00° 3; Hawkes Bay 1Jul99°; Port au 
Choix 20Jul99° 0; Squid Cove road 19Jul98°°9; St. John’s 
2Apr00° ; 

Clubiona furcata Emerton, 1919 [H] 

Holarctic. Mixed coniferous woods; litter under shrubs and 


herbs in marshy places. 
Main River west OOJul98° . 


258 


Clubiona johnsoni Gertsch, 1941 
Nearctic. Mixed coniferous woods. 
TNNP 15Aug80¢ (CNC). 


Clubiona kastoni Gertsch, 1941 
Nearctic. Mixed coniferous woods. 
Port au Choix 3-11Aug00°°¢. 


Clubiona kulczynskii Lessert, 1905 [H] 

Holarctic. Mixed coniferous woods; herbs in clearings in 
woods; Empetrum barrens. 

Cape Raven 15Aug99°; Gander 00Aug98°°; Main River 
west 00Jul98°S; Port au Choix 13Jul99°. 


Clubiona mixta Emerton, 1890 [H] - 
Nearctic. Shrubs and herbs. 
Twillingate Island 8Jul51° (FMNH). 


Clubiona moesta Banks, 1896 
Holarctic. Mixed coniferous woods. 
Pasadena 14Aug84° (CNC). 


Clubiona norvegica Strand, 1900 [H] 

Holarctic. Back of beaches; sand dunes; barren, rocky areas 
near the coast. Common in the west. 

Big Brook 14Jul01°°°; Conche 27Jul04°; New Ferolle 
18Aug00°°C0; Savage Cove 30Jun03°°S; St Pauls 
25Jul97° So. 


Clubiona obesa Hentz, 1847 [H] 

Nearctic. Shrubs and herbs. 

Deer Lake 30May51° (FMNH); Spruce Brook (Georges 
Lake) 8Jul49° (FMNH). 


Clubiona opeongo Edwards, 1958 
Nearctic. Exposed rocky barrens. 
Eddies Cove East 21Jun00¢. 


Clubiona riparia L. Koch, 1866 [H] 

Holarctic. Mixed coniferous woods; herbs in clearings in 
woods; vegetation at back of beaches. 

Bakers Brook estuary 26Aug98°; Green Gardens 5Jul979; 
Norris Point 20JunO00°; Stanford River 28Jul99° . 


Clubiona trivialis C. L. Koch, 1843 

Holarctic. Mixed coniferous woods; herbs in clearings in 
woods; mixed vegetation at back of beaches. 

Blackhead (St. John’s) 16Oct0O1°; Burnt Cape 4Aug98°; 
Gander 00Aug98° 3; Lobster Cove 23Jul9799; Norris Point 
27Jun00° . 


CYBAEIDAE (1 sp.) 

Cybaeota calcarata (Emerton, 1911) 
Nearctic. Litter of mixed coniferous woods. 
Norris Point 22Jul00° °. 


DICTYNIDAE (9 spp.) 

Argenna obesa Emerton, 1911 [H] 

Nearctic. Wetlands; river banks; clearings in woods. 

Come by Chance? (CNC); Grand Bank 3Aug51° ° (FMNH). 


Dictyna alaskae Chamberlin & Ivie, 1947 
Holarctic. Mixed coniferous woods; shrubs and herbs. 
Hampden 18-20Jun77° °? 99 (CNC). 


Dictyna arundinacea (Linnaeus, 1758) 

Holarctic. Mixed coniferous woods; shrubs and herbs. 

Bay d’Espoir 11Jul85°¢¢ (CNC); Hampden 12Jul77° 
(CNC); TNNP 7Jul87° (CNC). 


Dictyna bostoniensis Emerton, 1888 [H] 
Nearctic. Mixed coniferous woods; shrubs and herbs. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Joe Glodes Pond 21 Aug49° co (FMNH); Pasadena ° (CNC); 
St. John’s 21Jul862 (CNC). 


Dictyna brevitarsus Emerton, 1915 [H] 
Nearctic. Mixed coniferous woods; shrubs and herbs. 
Stag Brook (GMNP)17Jul99°¢. 


Dictyna minuta Emerton, 1888 

Nearctic. Mixed coniferous woods. 

Berry Head Pond (GMNP) 18Jul99°; Gadds Harbour 
27Jul97S (CNC); Norris Point 27Jul98° (and CNC). 


Emblyna annulipes (Blackwall, 1846) [H; Dictyna annulipes 
Blackwall] 

Holarctic. Mixed coniferous woods. 

Grandy Brook 2Jun49° (FMNH); Hampden 5- 29Jun86° ? 
SG (CNC); Pasadena 14Jun84° (CNC); Portland Creek 
OOJun749 (CNC). 


Emblyna phylax (Gertsch & Ivie, 1936) [H; Dictyna phylax 
Gertsch & Ivie] 

Nearctic. Mixed coniferous woods. 

Gander OOJul98¢. 


Iviella sp. 

Not attributed to species; probably new; to be described 
elsewhere. 

Nearctic. Only known from exposed sub-arctic Empetrum 
barrens on the Northern Peninsula. 

Burnt Cape 00Aug03° 9°; Killdevil Mountain 17Aug99°¢; 
Point Riche 17Aug98° °0°; Table Point 30Sep98¢. 


GNAPHOSIDAE (14 spp.) 

Drassodes neglectus (Keyserling, 1887) [H] 

Holarctic. Under stones on talus slopes; rocky barren areas; 
among stones at the back of beaches. 

Conche 16Jul03°°; Burnt Cape OOJul03°°9°; Killdevil 
Mountain 8Aug94¢; Point Riche 17Jul00°°; Port au Choix | 
4Jul039. 


Gnaphosa brumalis Thorell, 1875 

Nearctic. Wetlands; Empetrum barrens near coast. 
Burnt Cape 00Aug03°; Port au Choix 2Aug96°; Woody | 
Point (Lookout Hills) 5Jun97°. 


Gnaphosa microps Holm, 1939 

Holarctic. Wetlands; Empetrum barrens. 

Big Brook 21Jun02°; Killdevil Mountain 17Aug99° °c0; 
Point Riche 1Aug99G; Sally’s Cove 27Jul99°; St Lunaire | 
OO0Aug002. | 


Gnaphosa muscorum (L. Koch, 1866) [H] 
Holarctic. Empetrum barrens; rocky areas. 
Burnt Cape 00Aug03° °9c. 


Gnaphosa parvula Banks, 1896 [H] 
Nearctic. Empetrum barrens; rocky areas. 
Cape Ray 1Jul96°. 


Haplodrassus hiemalis (Emerton, 1909) [H] 

Holarctic. Empetrum barrens; rocky areas; peatlands. 
New Ferolle 13Jul04°; Point Riche 2Aug99° °; Table 
Point 30Sep98° °. 


Haplodrassus signifer (C. L. Koch, 1839) [H] 
Holarctic. Litter of shrubs and mixed coniferous woods. 
Burnt Cape 0O0Aug039¢. 


Micaria aenea Thorell, 1871 

Holarctic. Litter of shrubs and mixed coniferous woods; 
peatlands. 

Main River west OOJul98¢. 


2005 


Micaria constricta Emerton, 1894 [H; M. longispina Emer- 
ton, 1911] 

Holarctic. Empetrum barrens; rocky areas; peatlands; mixed 
coniferous woods. 

Base of Killdevil Mountain 13Jul49° (FMNH). 


Micaria pulicaria (Sundevall, 1831) [H] 

Holarctic. Empetrum barrens; rocky areas; peatlands; mixed 
coniferous woods. 

Gadds Harbour 11Aug99°; Point Riche 20Aug02°; Shal- 
low Bay 14Aug97°; Tablelands 18Aug99° ; Western Brook 
estuary 24Aug03°°. 


Micaria rossica Thorell, 1875 

Holarctic. Exposed sub-arctic Empetrum barrens on the 
Northern Peninsula. 

Burnt Cape 00Aug03°. 


Orodrassus canadensis Platnick & Shadab, 1975 [H; O. 
vastus Chamberlin & Ivie, 1922] 

Nearctic. Mixed coniferous woods. 

Doctors Brook 31Jul49° (FMNH): Eddies Cove West 
2Aug49? 2d (FMNH). 


Zelotes fratris Chamberlin, 1920 [H; Z. subterraneus (C. L. 
Koch, 1833)] 

Holarctic. Sand-dunes; rocky coastal areas; shrubby areas on 
lower slopes of mountains; abandoned meadows. Common. 
Badger 24Jun80°; Conche 27Jul04°; St. John’s OOJun952 
2°30; Tablelands 18Aug99° °°; Western Brook estuary 
25Jul972 PSS, 


Zelotes sula Lowrie and Gertsch, 1955 
Holarctic. Edge of peatlands; coastal Empetrum barrens. 
Burnt Cape 00Aug03° 29; Point Riche 19Aug00°o°. 


HAHNIIDAE (6 spp.) 

Antistea brunnea (Emerton, 1909) [H] 

Nearctic. Peatlands; wet parts of mixed coniferous woods. 
Bakers Brook estuary 9Aug99G; Berry Head Pond (GMNP) 
31Jul00S; Main River west OOJul98°°; Rocky Harbour 
23Jul-20Aug99°2 ° . 


Cryphoeca montana Emerton, 1909 [H; Agelenidae] 
Nearctic. Mixed coniferous woods; under bark; shrubby 
areas at lower elevations of mountains. 

Berry Head Pond (GMNP) 18Jul99° ; Gander 00Aug 980; 
| Port au Choix 28Jun99¢; Tablelands 21Jul99¢; TNNP 
} 1iJun93°. 


Hahnia cinerea Emerton, 1890 [H] 
| Nearctic. Peatlands and bogs. 
Burnt Cape 00Aug03° 2°90; Rocky Harbour 2Jul98¢. 


Hahnia glacialis Sérensen, 1898 

| Holarctic. Coastal and alpine Empetrum barrens; mixed con- 
iferous woods. 

Big Brook 16Jul01° °S; Conche 16Jul03° ; Nameless Cove 
16Aug01°°9°; Point Riche 15Jul00°°; Savage Cove 
} 14Jul01° eda, 


| Neoantistea agilis (Keyserling, 1887) [H] 

| Nearctic. Lower slopes of mountains under rocks; litter under 
| shrubs. 

| Tablelands 18Aug99¢¢. 


} Neoantistea magna (Keyserling, 1887) [H; N. riparia radula 
| (Emerton, 1890)] 

Nearctic. Mixed coniferous woods; coastal and alpine Empet- 
rum barrens; back of beaches; peatlands and bogs. 

Bakers Brook estuary 26Aug989; Big Brook 25Jun02° °; 


PICKAVANCE AND DONDALE: SPIDERS of NEWFOUNDLAND 


259 


Hawkes Bay 9Jul04°;; base of Killdevil Mountain 17Aug99° ¢ ; 
Sally’s Cove 10Aug99°c. 


LINYPHIDAE (191 spp.) 
Note: we follow Buckle et al. (2001) for linyphiid nomen- 
clature. 


Agyneta allosubtilis Loksa, 1965 [H; Argyneta decora (O. 
Pickard-Cambridge, 1871)] 

Holarctic. Mixed coniferous woods; shrubby areas. 

Burnt Cape 0O0Aug03° 2? 9; Killdevil Mountain 20Jul00c¢. 


Agyneta amersaxatilis Saaristo & Koponen, 1998 

Nearctic. Exposed coastal Empetrum barrens; rocky areas on 
exposed coasts. 

Point Riche 24Jul00¢9 (and CNC); Port au Choix 5Jul04¢. 


Agyneta dynica Saaristo & Koponen, 1998 [H; “Meioneta sp. 
pr rurestris’; this may be A. dynica] 

Nearctic. Exposed coastal and alpine Empetrum barrens; mixed 
coniferous woods. 

Cape Raven 15Aug99¢; Point Riche 24Jul00°¢. 


Agyneta fabra (Keyserling, 1886) 
Nearctic. Mixed coniferous woods. 
Gander 00Aug98¢; Corner Brook 10Aug94° . 


Agyneta jacksoni (Braendegaard, 1937) 
Nearctic. Mixed coniferous woods. 
Lewis Hills 21Aug86° (CNC). 


Agyneta olivacea (Emerton, 1882) [H; Argyneta cauta (O. 
P.-Cambridge, 1902)] 

Holarctic. Mixed coniferous woods. Common. 

Burnt Cape 00Aug03° °9°°; Gander 1Jul78°; Glide Lake 
OO0Jul94°2 3; Savage Cove 14Jul019¢; TNNP 10Jun93° °c. 


Agyneta simplex (Emerton, 1926) 

Nearctic. Exposed coastal Empetrum barrens. 

New Ferolle 13Jul04°; Point Riche 19Aug00° °; Burnt Cape 
O0Aug03? 2 dc. 


Agyneta sp. 

Not attributed to species (similar to CNC #6) 

Mixed coniferous woods. 

Main River west OOJul98°; Bakers Brook estuary 3Jul98°. 


Allomengea dentisetis (Grube, 1861) [H; Helophora ontari- 
ensis (Emerton, 1925)] 

Holarctic. Exposed coastal Empetrum barrens. 

New Ferolle 18Aug00° °9; Sally’s Cove 22Aug00° 9; St. 
Lunaire 15Aug00°. 


Aphileta misera (O. Pickard-Cambridge, 1882) [H; Eulaira 
concava Chamberlin & Ivie, 1945] 

Holarctic. Moist coniferous litter; coastal Empetrum barrens. 
Brunette Island ¢ (CNC?). 


Baryphyma kulczynskii (Eskov, 1979) 
Holarctic. Mixed coniferous woods. 
Pasadena 28May86°, 16May88° oo (CNC). 


Bathyphantes brevipes (Emerton, 1917) [H] 

Nearctic. Mixed coniferous woods; stunted coniferous stands 
near coast; shrubs in road cuttings. 

Big Brook 19Jun02°°9°; Nameless Cove 16Aug01°°; 
Point Riche 1 Aug99¢; Port au Cho1x 13 Jul99°; Rocky Har- 
bour 30Jul97¢. 


Bathyphantes brevis (Emerton, 1911) [H] 
Nearctic. Litter around pools on Empetrum barrens. 
Badger 22-25Jun51° (FMNH); Cow Head 8Aug49° (FMNH); 


260 


Deer Lake 30May51° (FMNH); South Branch (Codroy) 
4Jul49° (FMNH). 


Bathyphantes canadensis (Emerton, 1882) 

Holarctic. Mixed coniferous woods. 

Georges Lake 13Sep88° (CNC); Pasadena 28May86? (CNC); 
Steady Brook 1 1Jun84° (CNC). 


Bathyphantes concolor (Wider in Reuss, 1834) [H] 
Holarctic. Mixed coniferous woods; thickets of alder and other 
shrubs; coastal Empetrum barrens; back of beaches; gardens 
and cultivated areas. Common. 

Bakers Brook estuary 8-27Aug00° 9; Mount Scio 00Aug959; 
Port au Choix 1 Aug99°'0; Stanford River 29Jul99¢; St. John’s 
2Apr00°¢C. 


Bathyphantes gracilis (Blackwall, 1841) [H] 

Holarctic. Peatlands and bogs. 

Berry Head Pond (GMNP) 19-31Jul00° °9°; Burnt Cape 
O0AugO3° Peds. 


Bathyphantes pallidus (Banks, 1892) [H; Bathyphantes sp. 
probably reprobus (Kulczynski, 1916)] 

Nearctic. Mixed coniferous woods; coastal Empetrum barrens. 
Bakers Brook estuary 8-27Aug00°°9°; Burnt Cape 
O00Aug03°; base of Killdevil Mountain 13Aug00° °c. 


Bathyphantes reprobus (Kulczyhski, 1916). 

Marusik et al. (1993) synonymised B. crosbyi (Emerton, 1919) 
under B. reprobus. CDD thinks the species are distinct and 
that Newfoundland specimens are B. crosbyi; see Buckle et 
al. 2001. 

Holarctic. Mixed coniferous woods. 

Main River west 00Jul98¢. 


Bathyphantes simillimus (L. Koch, 1879) 

Holarctic. Mixed coniferous woods. 

Berry Head Pond (GMNP) 31Jul-15Aug00°; Glide Lake 
OOJul94°; Main River west OOJul98°°; Stanford River 
29Jul99C. 


Carorita limnaea (Crosby & Bishop, 1927) 
Holarctic. Peatlands; mosses in mixed coniferous woods. 
Main River west 23Jul00°¢. 


Centromerita bicolor (Blackwall, 1833) [H; Centromerus 
bicolor (Blackwall)] 

Palaearctic; introduced. Mixed coniferous woods; coastal 
Empetrum barrens. 

Blackhead (St. John’s) 7Oct-12Nov? °oc. 


Centromerus cornupalpis (O. Pickard-Cambridge, 1875) [H] 
Nearctic. Mixed coniferous woods. 
Grand Bruit 13Jun49° (FMNH). 


Centromerus denticulatus (Emerton, 1909) 
Nearctic. Mixed coniferous woods; sphagnum bogs. 
Rocky Harbour (Bottom Brook bog) 6Aug98°. 


Centromerus furcatus (Emerton, 1882) 

Nearctic. Mixed coniferous woods. 

Berry Head Pond (GMNP) 26Jul-14Aug00°°; Cook’s Pond 
4-10Jul940; Glide Lake 00Jul94° ; Port au Choix 13Jul99oC. 


Centromerus latidens (Emerton, 1882) [H] 

Nearctic. Mixed coniferous woods. 

Burgeo, Cow Head, Gambo, Grand Bruit, Millertown: 11- 
20Jun49° ; 1-31Jul49 °°; 1-31 Aug49° ° (FMNH_?). Dates not 
connected to localities; year assumed 1949 from context 
(Hackman 1954). Not found by CDD in Hackman’s collec- 


THE CANADIAN FIELD-NATURALIST 


Volt 


tions in FMNH. This record may be erroneous but is retained 
here pending clarification. 


Centromerus longibulbus (Emerton, 1882) [H] 

Nearctic. Mixed coniferous woods; sphagnum bogs. 

Berry Head Pond (GMNP) 31Jul-15Aug00°; Main River 
west OOJul98¢; Port au Choix 8Jul00¢; TNNP 10Jun93¢. 


Centromerus persolutus (O. Pickard-Cambridge, 1875) [H] 
Nearctic. Mixed coniferous woods. 
Corner Brook 10Aug94°; Glide Lake 00Jul94¢. 


Centromerus sylvaticus (Blackwall, 1841) [H] Holarctic. 
Mixed coniferous woods. 

Cow Head 10Aug49° (FMNH?); Grandy Brook 24Jun49° 
(FMNH?). Not found by CDD in Hackman’s collections in 
FMNH. This record may be erroneous but is retained here 
pending clarification. 


Ceraticelus alticeps (Fox, 1891) 
Nearctic. Mixed coniferous woods. 
Corner Brook 17Aug84° (CNC). 


Ceraticelus atriceps (O. Pickard-Cambridge, 1874) [H] 
Nearctic. Mixed coniferous woods. Common. 

Bakers Brook estuary 15Jul99°; Big Brook 24Jun02° °; 
Burnt Cape 4Aug98¢; Norris Point 28Jul98° 9; Port au Choix 
12Jul99° 


Ceraticelus bulbosus (Emerton, 1882) 

Holarctic. Mixed coniferous woods. 

Bay d’Espoir 11Jul85° (CNC); Brunette Island 18May85? 
(CNC); Eddies Cove (East or West not specified) 11 Aug76? 
(CNC); Portland Creek 8Sep85° ° (CNC). 


Ceraticelus fissiceps (O. Pickard-Cambridge, 1874) [H] 
Nearctic. Mixed coniferous woods. Common. 

Burnt Cape 00Aug03°°; Cow Head 7Aug97°; Gander 
28Jul98¢; Port au Choix 12 Jul99¢; Stanford River 28Jul99¢. 


Ceraticelus laetabilis (O. Pickard-Cambridge, 1874) [H] 
Nearctic. Mixed coniferous woods. 
Glide Lake 00Jul94¢. 


Ceraticelus laetus (O. Pickard-Cambridge, 1874) 
Nearctic. Sphagnum bogs; mixed coniferous woods. 
Rocky Harbour (Bottom Brook bog) 20Aug99°. 


Ceraticelus minutus (Emerton, 1882) 

Nearctic. Mixed coniferous woods. 

Gambo Pond 29Oct85° (CNC); Highlands River 14Aug84? | 
2°35 (CNC); New Bay Pond 14Aug84° (CNC); Pasadena | 
19Jul84¢ (CNC); Triton Brook 290ct85° So (CNC). 


Ceraticelus similis (Banks, 1892) [H] 
Nearctic. Empetrum barrens; mixed coniferous woods. 
Shallow Bay 9Aug98¢. 


Ceratinella brunnea Emerton, 1882 [H] 
Nearctic. Mixed coniferous woods; Empetrum barrens; sphag- — 
num bogs. Common. 
Big Brook 16Jul01°; Blackhead (St. John’s) 21Apr99°; Burnt — 
Cape 00Aug03°9; Norris Point 22Jul00¢; Port au Choix | 
1Aug99°. 


Ceratinella ornatula (Crosby & Bishop, 1925) 
Nearctic. Mixed coniferous woods. 
Hampden 29Jun77° (CNC). 


Ceratinella parvula (Fox, 1891) 
Nearctic. Mixed coniferous woods; rocky barrens. 
Eddies Cove East 21Jun00° ; Main River west O0Jul98¢. 


2005 


Ceratinops annulipes (Banks, 1892) 
Nearctic. Sphagnum bogs. 
Rocky Harbour (Bottom Brook bog) 13Aug98°. 


Ceratinopsis nigriceps Emerton, 1882 
Nearctic. Mixed coniferous woods. 
Badger 30Jul57° (CNC); Gander 14Jul789 (CNC). 


Ceratinopsis nigripalpis Emerton, 1882 
Nearctic. Mixed coniferous woods. 
Gallows Cove 2Nov97°2. 


Cheniseo sphagnicultor Bishop & Crosby, 1935 
Nearctic. Sphagnum bogs. 
Rocky Harbour (Bottom Brook bog) 13Aug99°. 


Dicymbium elongatum (Emerton, 1882) 
Nearctic. Mixed coniferous woods. 
Main River west 24Jul00°. 


Dicymbium nigrum (Blackwall, 1834) 

Palaearctic; introduced. Mixed coniferous woods; shrubs in 
abandoned meadows; shrubs and trees in gardens. These are 
the only known North American records. 

Gallows Cove 2Nov97°C (and CNC); St. John’s (Forest 
Avenue) 18Apr99°; St. John’s (University) 26Sep99°. 


Dietrichia hesperia Crosby & Bishop, 1933 [H] 

Nearctic. Mixed coniferous woods. 

Stephenville 290ct473 (AMNH7?). This record is based on a 
single male collected by Robert Traub and lent to Hackman 
by Dr. W. J. Gertsch. Hackman (1954) discussed whether the 
Newfoundland specimen was conspecific with a paratype 
(pages 17, 18) and put a question mark by this species in his 
checklist (page 6). It is assumed that Hackman returned the 
specimen to the AMNH. This record may be erroneous but 
is retained here pending clarification. 


Diplocentria bidentata (Emerton, 1882) [H] 

Holarctic. Mixed coniferous woods; shrubs; sphagnum bogs; 
Empetrum barrens; abandoned meadows; gardens. Common. 
Berry Head Pond (GMNP) 4Aug99¢; Big Brook 19Jun02° ; 
Lobster Cove 22Jul99¢; Norris Point 28Jul98° ; Point Riche 
1Aug99¢c. 


Diplocentria rectangulata (Emerton, 1915) 
Holarctic. Mixed coniferous woods. 
Main River west OOJul98°¢. 


Diplocephalus cristatus (Blackwall, 1833) [H] 
Holarctic. Mixed coniferous woods. 
Port au Choix 13Jul99¢c. 


Diplocephalus subrostratus (O. Pickard-Cambridge, 1873) 
[H; Diplocephalus cuneatus (Emerton, 1909)] 

Holarctic. Mixed coniferous woods. 

Lloyds River 19Jun84° °¢ (CNC); Millertown 20Aug84° 
3 (CNC); Pasadena 23May88° ° (CNC); Point Leamington 
} 24Sep84? (CNC). 


| Dismodicus alticeps Chamberlin & Ivie, 1947 [H] 
| Holarctic. Mixed coniferous woods. 
| Noel Paul’s Brook 8Jul77° (CNC). 


| Dismodicus decemoculatus (Emerton, 1882) [H; Dismodicus 
| bifrons decemoculatus (Emerton)]| 
Nearctic. Coastal Empetrum barrens; litter and rocks at back 


| of beaches. 


| Big Brook 22Jun02°°9¢; Port au Choix 5Jul04¢; Sally’s 
Cove 21Jul00°; St. Lunaire 15Aug00°°; Watts Point 
29Jul98° . 


PICKAVANCE AND DONDALE: SPIDERS Of NEWFOUNDLAND 


261 


Drapetisca alteranda Chamberlin, 1909 
Nearctic. Mixed coniferous woods. 
St. John’s 0000222 (CNC). 


Entelecara sombra (Chamberlin & Ivie, 1947) 

Holarctic. Empetrum barrens at higher elevations and on tops 
of mountains; rocky coastal barrens in exposed places. 

Big Brook 25Jun02° ° ; Conche 27Jul04° ; Highlands of St. 
John 14Aug002° ; Killdevil Mountain 26Jul99° 3; Point Riche 
24Jul002 S. 


Eperigone bryantae Ivie & Barrows, 1935 
Nearctic. Mixed coniferous woods. 
Renews 28Jul72° (CNC). 


Eperigone entomologica (Emerton, 1911) 

Nearctic. Sphagnum bogs; mixed coniferous woods. 

Bakers Brook estuary 14Jul98°; Rocky Harbour (Bottom 
Brook bog) 12Aug98°°; Main River west 00Jul98¢. 


Eperigone index (Emerton, 1914) 
Nearctic. Sphagnum bogs; mixed coniferous woods. 
Fermeuse 17Oct85° (CNC). 


Eperigone maculata (Banks, 1892) [H] 
Holarctic. Mixed coniferous woods. 
Gander 17Jul78°°; Little Grand Lake 27Jul94¢. 


Eperigone trilobata (Emerton, 1882) [H] 
Holarctic. Coastal Empetrum barrens; mixed coniferous woods. 
Blackhead (St. John’s) 16OctO1¢. 


Eperigone undulata (Emerton, 1914) [H; Eperigone contorta 
(Emerton, 1882)] 

Nearctic. Coastal Empetrum barrens; mixed coniferous woods. 
Bakers Brook estuary 9Aug99°; Main River west 00Jul98¢. 


Eridantes utibilis Crosby and Bishop, 1933 [H; Scylaceus 
obtusus Emerton, 1915] 

Nearctic. Mixed coniferous woods. 

Corner Brook 25Jul92° ; Squid Cove road 19Jul98° ; Port au 
Choix 24Jun00°. 


Erigone aletris (Crosby & Bishop, 1928) [H; Erigone ephala 
Crosby & Bishop, 1928] 

Holarctic. Mixed coniferous woods; sphagnum bogs; litter, 
stones and low herbs at back of beaches; Empetrum barrens. 
Bakers Brook estuary 13Sep97°; Cape Freels 12Jul01° °; 
New Ferolle 10Aug00° 9°30; Savage Cove 30Jun03° °; Seal 
Cove (Bonne Bay) 14Jul98¢. 


Erigone atra Blackwall, 1833 [H] 
Holarctic. Mixed coniferous woods. 
Pasadena 28May869° (CNC). 


Erigone blaesa Crosby & Bishop, 1928 [H] 

Nearctic. Litter at back of beaches; litter around lakes; sand 
dunes. 

Brunette Island 14Jul94°; Cape Freels 12 Jul01°°9°; 
Western Brook Pond? °. 


Erigone dentigera (O.Pickard-Cambridge, 1874) [H] 
Holarctic. Mixed coniferous woods; abandoned meadows; 
sphagnum bogs. 

Big Brook 24Jun02°; Gadds Harbour 28Jul99¢ ; Main River 
west 29Jul00¢. 


Erigone dentipalpis (Wider in Reuss, 1834) 

Palaearctic; introduced. Coastal Empetrum barrens; mixed 
coniferous woods; gardens. 

Blackhead (St. John’s) 25Jul03¢; Main River west 25Jul00¢c; 
St. John’s 40ct02¢. 


262 


Erigone ephala Crosby & Bishop, 1928 [H] 

Nearctic. Mixed coniferous woods; back of beaches. 

Berry Hill (GMNP) 29Jul76¢ (CNC); Brunette Island 7- 
8May86° (CNC); Chance Cove 25Jun85° 9°30 (CNC); East- 
port 7Aug769 (CNC); Stephenville 30Mar87¢ (CNC). Buckle 
et al. (2001) say “probably = E. aletris”. Retained here pend- 
ing clarification. 


Erigone mentasta Chamberlin and Ivie, 1947 [H; female 
Hilaira mentasta (Chamberlin and Ivie); male a misidentifi- 
cation of Sciastes dubius (Hackman, 1954)] 

Nearctic. Mixed coniferous woods. 

Lomond 14Jul49° (FMNH). 


Erigone whymperi O. Pickard-Cambridge, 1877 [H] 
Nearctic. Mixed coniferous woods; Empetrum barrens. 
Cooks Harbour 16Jul49° (FMNH); Doctors Hill (= Highlands 
of St. John, Port au Choix, not Doctor’s Hill, Port Blandford) 
29Jun49° (FMNH); Flowers Cove 25Jul49° (FMNH); Hare 
Bay 14Jul49° (FMNH); Waterford Bridge 5Jun49° (FMNH). 
These records of an essentially arctic species (Buckle et al. 
2001) seem surprising. Hackman (1954, pages 18, 58) adds 
a question mark to his identification. Retained here pending 
clarification. 


Estrandia grandaeva (Keyserling, 1886) [H] 

Holarctic. Mixed coniferous woods. 

Berry Head Pond (GMNP) 24Jul99¢; Lomond 12Aug93° ; 
Port au Choix 12Jul9990; Shallow Bay 13Jul97° ° (and CNC); 
Western Brook Pond 5Jul99° ° (and CNC). 


Eulaira microtarsus (Emerton, 1882) [H] 

Nearctic. Mixed coniferous woods. 

Corner Brook 24Jul92°; Glide Lake OOJul94° °°; Main 
River west OOJul98°c. 


Floricomus rostratus (Emerton, 1882) 
Nearctic. Mixed coniferous woods. 
Main River west OO0Jul98°. 


Glyphesis scopulifer (Emerton, 1882) [H; Tapinocyba scop- 
uliferum (Emerton) | 

Nearctic. Mixed coniferous woods; edges of waterways. 
Cow Head 8Jul492° (FMNH). 


Gnathonaroides pedalis (Emerton, 1923) 
Nearctic. Mixed coniferous woods. 
Corner Brook 25Jul92° ; Corner Brook 14Jun93¢. 


Gonatium crassipalpum Bryant, 1933 

Nearctic. Coastal rocky barrens; Empetrum barrens; sphagnum 
bogs. 

Berry Head Pond (GMNP) 19Jul00° ; Logy Bay 20Nov03° ; 
Point Riche 13Jul99¢; St. Lunaire 15Aug00°; Table Point 
30Sep98¢C. 


Grammonota angusta Dondale, 1959 [H; Grammonota pic- 
tilis (O. Pickard-Cambridge, 1875)] 

Nearctic. Mixed coniferous woods; low vegetation in clearings 
in woods. Common. 

Gadds Harbour 27Jul97°°; Gallows Cove 10Aug99° °; 
Gander 28Jul989°; Port au Choix 8Jul00° ; Stuckless Pond 
20Jul93¢c. 


Grammonota capitata Emerton, 1924 [H; Grammonota pic- 
tilis (O. Pickard-Cambridge, 1875) in part] 

Nearctic. Mixed coniferous woods. 

South Branch (Codroy) 3Jul49° (FMNH); Spruce Brook River 
8-9Jul49° 2° (FMNH); Victoria Lake 13Jun51° (FMNH). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Grammonota gigas (Banks, 1896) [H] 

Nearctic. Mixed coniferous woods; sphagnum bogs. 

Bakers Brook estuary 14Jul98° °; Rocky Harbour (Bottom 
Brook bog) 13Aug98° ; Main River west 00Jul98° 3; Mount 
Scio 0O0Aug95°. 


Grammonota maritima Emerton, 1925 [H] 
Nearctic. Empetrum barrens; rocky coastal barrens. 
Burnt Cape 00Aug03°¢o. 


Grammonota vittata Barrows, 1919 
Nearctic. Low vegetation in and around sphagnum bogs. 
Rocky Harbour (Bottom Brook bog) 16Jul99¢. 


Halorates alascensis (Banks, 1900) [H; Collinsia clypiella 
(Chamberlin, 1920)] 

Nearctic. Salt marshes; wet Empetrum barrens. 

Cow Head 10Aug49° (FMNH?); Deer Lake 30May51° 2? do 
(FMNH?). Not found by CDD in Hackman’s collections in 
FMNH. This record may be erroneous but is retained here 
pending clarification. 


Halorates holmgrenii (Thorell, 1871) 
Holarctic. Exposed coastal barrens. 
Big Brook 19Jun02°. 


Halorates palmeni (Hackman, 1954) [H; Collinsia palmeni 
Hackman] 

Known only from a single female reported by Hackman (1954) 
Nearctic. River bank. 

South Branch (Codroy) 2Jul49° (CNC). 

Note: CDD and J. Redner examined this specimen and think 
it is Oedothorax maximus (Emerton); this question will be 
dealt with elsewhere. 


Halorates pertinens (O. Pickard-Cambridge, 1875) [H; Col- 
linsia pertinens O. Pickard-Cambridge] 

Nearctic. Among stones and litter at the back of beaches. 
Big Brook 18Jun02° 9°; Gadds Harbour 11 Aug99°. 


Helophora insignis (Blackwall, 1841) [H] 
Holarctic. Mixed coniferous woods. 
Bakers Brook estuary 27Aug00°. 


Hilaira canaliculata (Emerton, 1915) [H; Hilaira aquilonia 
Hackman, 1954] 

Holarctic. In litter and moss in thickets of shrubs. 

Stanford River 11 Aug49? ° (CNC). 


Hilaira herniosa (Thorell, 1875) [H] 
Holarctic. Exposed coastal Empetrum barrens. 
Raleigh 17Jul48° (FMNH). 


Hybauchenidium gibbosum (S@rensen, 1898) [H; Hybocop- 
tus denticulatus (Emerton, 1915)] 

Nearctic. Mixed coniferous woods. 

Main River west OOJun98° C. 


Hypomma marxii (Keyserling, 1886) [H] 

Nearctic. Empetrum barrens; sphagnum bogs. 

Cow Head 11Aug49° (FMNH); South Branch (Codroy) 
2Jul49° (FMNH); Victoria Lake 11Jun51° ° (FMNH). 


Islandiana flaveola (Banks, 1892) 
Nearctic. Mixed coniferous woods. 
Main River west 31Jul00¢C. 


Islandiana holmi Ivie, 1965 [H; Islandiana alata (Emerton, 
1919)| 

Nearctic. Empetrum and rocky barrens at the coast and 
higher altitudes. 

Bartletts Harbour 12Jul04°; Port au Choix 5 Jul04°. 


2005 


Islandiana princeps Braendegaard, 1932 [H; /slandiana 
alata (Emerton, 1919)] 

Nearctic. Sphagnum bogs; Empetrum barrens. 

Big Brook 22Jun02¢; Burnt Cape 0D0Aug03° 3; Sally’s Cove 
bog 27Jul99¢. 


Kaestneria pullata (O. Pickard-Cambridge, 1863) [H; Bathy- 
phantes pullatus (O. Pickard-Cambridge)] 

Holarctic. Shrubs and herbs in mixed coniferous woods; shrubs 
and herbs in road cuttings. 

Bellburns 8Jul97°; Port au Choix 20Jul99°; Sally’s Cove 
31Jul-15Aug00°; Shallow Bay 9Aug98° ; TNNP 10Jun93¢. 


Kaestneria rufula (Hackman, 1954) [H; Bathyphantes rufulus 
Hackman] 

Nearctic. Sphagnum bogs. 

Bakers Brook estuary 3Jul98°. 


Lepthyphantes alpinus (Emerton, 1882) [H] 

Holarctic. Mixed coniferous woods. 

Gander 20Jul98°9°; Killdevil Mountain 20Jul00° °o°o; 
Norris Point 27Jul98°; Point Riche 1Aug99° °C; Port au 
Choix 1Aug99°°; Stanford River 29Jul99° co. 


Lepthyphantes bihamatus (Emerton, 1882) [H] 

Nearctic. Mixed coniferous woods; low herbs in clearings in 
woods. 

Berry Head Pond (GMNP) 4Aug99¢; Lobster Cove 22Jul99¢ ; 
Norris Point 14Aug00°; Port au Choix 12Jul99°; Shallow 
Bay 13Jul97¢. 


Lepthyphantes calcaratus (Emerton, 1909) [H] 

Nearctic. Mixed coniferous woods; stunted trees near coast; 
sea-cliffs. 

Berry Head Pond (GMNP) 24Jul99°; Green Gardens 
O0Aug929; base of Killdevil Mountain 20Jul002; Lobster 
Cove 9Aug990'G; Point Riche 24Jul98¢. 


Lepthyphantes complicatus (Emerton, 1882) [H; Lepthy- 
phantes umbraticola (Keyserling, 1886)] 

Holarctic. Mixed coniferous woods; coastal rocky and Empet- 
rum barrens. 

Big Brook 19Jun99°°; Nameless Cove 16Aug01°; Point 
Riche 13Jul999; Port au Choix 1Aug99¢; Savage Cove 
23Aug98°. 


Lepthyphantes cracens Zorsch, 1937. [H; Lepthypantes nigri- 
ventris (L. Koch, 1879)] 

Nearctic. Mixed coniferous woods. 

Previously reported only from New York State (Buckle et al. 
2001) and Quebec (Paquin & Dupérré 2003 as Tenuiphantes) 
on the basis of males. Both males and females have been 
found in Newfoundland, and the female has been described 
elsewhere (Pickavance 2004). The female is similar to L. 
nigriventris (L. Koch, 1879), and likely Hackman’s (1954) 
| report of a single female of this Palaearctic species from New- 
| foundland was a misidentification of a specimen of L. 
| cracens. We feel that L. nigriventris probably does not occur 
| in North America. 

| Port au Choix 2-11Aug00° °? Sc, 20Aug00° ° 3S (and CNC). 


Lepthyphantes duplicatus (Emerton, 1913) [H; Lepthyphantes 
triramus Chamberlin & Ivie, 1947] 
_ Nearctic. Mixed coniferous woods. 
Bakers Brook estuary 8-27Aug°? 3; Little Grand Lake 6Aug 
| 94°: Norris Point 1S5Aug00° ; Port au Choix 8- 24Jul00¢. 


Lepthyphantes intricatus (Emerton, 1911) 
| Nearctic. Mixed coniferous woods. 
| Main River west 00Jul98¢. 


PICKAVANCE AND DONDALE: SPIDERS Of NEWFOUNDLAND 


263 


Lepthyphantes leprosus (Ohlert, 1865) [H] 

Palaearctic; introduced. Mixed coniferous woods; sea cliffs; 
houses and outbuildings. 

Blackhead (St. John’s) 12Nov02°; Mt. Scio 8Oct99° ; Port au 
Choix 1Oct98¢; St. John’s 2Nov02¢; Tuckers Head 30ct98°. 


Lepthyphantes tenuis (Blackwall, 1852) 

Palaearctic; introduced. Coastal Empetrum barrens near hu- 
man settlements. 

Blackhead (St. John’s) 12Nov029°; Logy Bay 16Sep03°. 


Lepthyphantes turbatrix (O. Pickard-Cambridge, 1877) [H; 
Lepthyphantes subalpinus (Emerton, 1882)]| 

Nearctic. Mixed coniferous woods; stones at the back of beach- 
es; sea cliffs; talus slopes. 

Boutitou 17Jul03°; Conche 16Jul03° °; Gander 26Jul98¢; 
Lomond 12Aug93¢; Tuckers Head 30Jul972 °°. 


Lepthyphantes washingtoni Zorsch, 1937 

Nearctic. Rocky or Empetrum barrens; mixed coniferous 
woods. 

Big Brook 20-25Jun03° ° ; Main River west OOJul002 ; Name- 
less Cove 16Aug01°; Point Riche 25Jun00° ; Port au Choix 
3-11Aug?s. 


Lepthyphantes zebra (Emerton, 1882) [H] 
Nearctic. Mixed coniferous woods. 
Logy Bay 6Oct02°. 


Lepthyphantes spp. 

Two unidentified species. In CNC as Lepthyphantes #2, Lep- 
thyphantes #7. 

Nearctic. Mixed coniferous woods. 

#2 Berry Head Pond (GMNP) 24Jul99° (and CNC); #7 Glide 
Lake 00Jul9499; Main River west 1Aug00° (and CNC). 


Lophomma sylvaticum (Emerton, 1913) [H; a specimen of 
this species labelled Entelecara is in Hackman’s material in 
FMNH, but is not recorded in his list of species] 

Nearctic. Mixed coniferous woods. 

Main River west OOJul98°2¢. 


Maro amplus Dondale & Buckle, 2001 

Nearctic. Mixed coniferous woods; shrubs and herbs in clear- 
ings in woods. 

Corner Brook 24Jul92¢; Main River west 17Jun989°; Mt. 
Scio OOAug95¢ (and CNC); Glide Lake 00Jul94¢¢. 


Maro nearcticus Dondale & Buckle, 2001 

Nearctic. Mixed coniferous woods. 

Little Grand Lake 15Jul92°, 13Jul93°°; Main River west 
27Jul002 ; Stanford River 29Jul99° . 


Maso sundevallii (Westring, 1851) [H] 
Holarctic. Mixed coniferous woods. 
Main River west OOJul98¢. 


Mecynargus paetulus (O. Pickard-Cambridge, 1875). 
Holarctic. Coastal subarctic barrens. 
Nameless Covel6Aug01°; Savage Cove 30Jun03°. 


Micrargus longitarsus (Emerton, 1882) 
Nearctic. Mixed coniferous woods. 
Corner Brook 7Aug93°¢. 


Microlinyphia mandibulata (Emerton, 1882) [H; Pusillia 
mandibulata Emerton] 

Nearctic. Coastal Empetrum barrens; moist abandoned 
meadows. 

Badger 24Jun80° ; Blackhead (St. John’s) 10Jun01¢; Logy 
Bay 6Aug01°; Main River west 14Aug00°. 


264 


Mythoplastoides exiguus (Banks, 1892) [H; Entelecara exigua 
(Banks)] 

Nearctic. Sphagnum bogs. 

Pushthrough 24Jun49° ° (FMNH?). Not found by CDD in 
Hackman’s collections in FMNH. This record may be erro- 
neous but is retained here pending clarification. 


Neriene clathrata (Sundevall, 1830) [H; Linyphia waldea 
Chamberlin & Ivie, 1943] 

Holarctic. Mixed coniferous woods; coastal abandoned mead- 
ows; shrubs in road cuttings. 

Bay Bulls 1May96°; Blackhead (St. John’s) 9Jun03¢ ; Gander 
28Jul982; Gros Morne Mountain 16Jun94°; Norris Point 
7Jul00¢. 


Neriene radiata (Walckenaer, 1841) [H; Linyphia marginata 
C. L. Koch, 1834] 

Holarctic. Mixed coniferous woods; sea cliffs; shrubs in road 
cuttings. 

Gander 28Jul989°; Rocky Harbour 30Jul97° ; Tuckers Head 
30Jul97° . 


Oedothorax trilobatus (Banks, 1896) 

Nearctic. Mixed coniferous woods; sphagnum bogs; rocky 
barrens. 

Bakers Brook estuary °; Berry Head Pond (GMNP) 22Aug 
00° °; Rocky Harbour (Bottom Brook bog) 23Jul99° ; Eddies 
Cove East 21Jun00° 3; Main River west 20Jul00¢. 


Oreonetides flavescens (Crosby, 1937) [H] 
Nearctic. Sphagnum bogs. 
Rocky Harbour (Bottom Brook bog) 5Aug98°. 


Oreonetides rectangulatus (Emerton, 1913) 

Nearctic. Sphagnum bogs. 

Berry Head Pond (GMNP) 15-22Aug00° ° 9°; Rocky Har- 
bour (Bottom Brook bog) 23Jul-20Aug99° 0; Sally’s Cove 
6-21Jul00s¢c. 


Oreonetides rotundus (Emerton, 1913) [H; Diplocentria 
corynetes Chamberlin & Ivie, 1945] 

Nearctic. Mixed coniferous woods. 

Gallows Cove 2Nov97° ; Glide Lake 0D0Aug94¢; Main River 
west OOJul98° ; Norris Point 28Jul98° So. 


Oreonetides vaginatus (Thorell, 1872) [H] 

Holarctic. Mixed coniferous woods. 

Glide Lake 00Aug96°°; Main River west OOJul98°; Port 
au Choix 8Jul00¢. 


Oreonetides spp. 

Two unidentified species. In CNC as Oreonetides #1 and Ore- 
onetides #2 

Nearctic. Mixed coniferous woods. 

#1 Norris Point 28Jul98° , 20Jun-13Jul00° ° ; #2 Glide Lake 
O0Aug96C. 


Oreophantes recurvatus (Emerton, 1913) 

Nearctic. Mixed coniferous woods. 

Bakers Brook estuary 8-27Aug00°°9; Berry Head Pond 
(GMNP) 26Jul00¢; Glide Lake 00Aug96¢°; Little Grand 
Lake 27Jul94°. 


Pelecopsis mengei (Simon, 1884) [H; Trichopterna mengei 
(Simon)} 

Holarctic. Mixed coniferous woods. 

Burnt Cape 0O0Aug03° ; Main River west 00Jul98°¢. 


Pityohyphantes costatus (Hentz, 1850) [H; Pityohyphantes 
costata Hentz] 
Nearctic. Mixed coniferous woods. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Deer Lake 3Jun51° (FMNH); Hare Bay 21Jun49° (FMNH); 
Lomond 14Jul49° (FMNH): Rencontre 17-19Jun49¢ 
(FMNH); Spruce Brook 8-9Jul49¢ (FMNH). 


Pityohyphantes limitaneus (Emerton, 1915) 

Nearctic. Mixed coniferous woods. 

Berry Head Pond (GMNP) 29Jun00° °90; Lomond 14Jul98° 
°; Millbrook (Rocky Harbour) 4Aug97? °. 


Pityohyphantes subarcticus Chamberlin & Ivie, 1943 [H; 
Pityohyphantes costata Hentz, 1850] 

Nearctic. Mixed coniferous woods; rocky barrens. 

Bakers Brook estuary 8-27Aug00°°; Eddies Cove East 
21Jun00° ; Gander 28Jul98° ° ; Main River west 28Jul00° ° ; 
Port au Choix 24Jul00¢. 


Pocadicnemis americana Millidge, 1976 

Nearctic. Mixed coniferous woods; rocky barrens; stones and 
litter at back of beaches. 

Bakers Brook estuary 6Aug99°; Burnt Cape 00Aug03°; 
Gander 28Jul98° 9; Little Grand Lake 22Aug94°; Mt. Scio 
O0AUg9SS. 


Poeciloneta furcata (Emerton, 1913) 
Nearctic. Mixed coniferous woods. 
Little Grand Lake 24Aug93¢. 


Poeciloneta sp. 

Unidentified species. NF 017 male and NF 108 female. 
Nearctic. Mixed coniferous woods. 

NFO17 Corner Brook 19Jul-17Aug939 (CNC); NF108 Cor- 
ner Brook 10-23Aug94? (CNC). 


Porrhomma gertschi Hackman, 1954 [H] 
Nearctic. Mixed coniferous woods. 
Known only from Newfoundland. 

Deer Lake 30May51°9 (CNC). 


Porrhomma terrestre (Emerton, 1882) 

Nearctic. Mixed coniferous woods. 

Berry Head Pond (GMNP) 16Aug99¢, 26Jul-4Aug00¢9; 
Corner Brook 14Jun-6Jul93°9, 19Jul-17Aug93°; base of 
Killdevil Mountain 20Jul00° . 


Porrhomma sp. 

Unidentified species: Porrhomma sp. #1 
Nearctic. Mixed coniferous woods. 

Glide Lake 00Jul94° ; Corner Brook 25Jul92°. 


Satilatlas gertschi Millidge, 1981 
Nearctic. Exposed coastal barrens. 
Burnt Cape 00Jul03¢. 


Satilatlas marxii Keyserling, 1886 

Holarctic. Mixed coniferous woods; Empetrum barrens. 
Burnt Cape O00Jul03° ° ; Eddies Cove East 21Jun00° ° ; Main 
River west OOJul98° . 


Sciastes dubius (Hackman, 1954) [H; Hilaira dubia Hackman] 
Holarctic. Mixed coniferous woods; damp moss. 

Grandy Brook 26Jun49° °? (FMNH); Rose Blanche 27Jun49° 
(FMNH); St. Barbe 26Jul49° (FMNH); Stanford River 
11Aug49? (CNC); Victoria Lake 11- 13Jun51° (FMNH). 


Sciastes truncatus (Emerton, 1882) [H] 

Nearctic. Mixed coniferous woods. 

Gander 28Jun98°; Glide Lake 00Jul949; Main River west 
OOJul989; Norris Point 27Jul98° , 26Jul00cS. 


Scirites pectinatus (Emerton, 1911) [H; Tapinocyba exigua 
Hackman, 1954] 


2005 


Nearctic. Mixed coniferous woods; birch stands. 
Millertown Junction 22Aug49° (CNC). 


Scironis tarsalis (Emerton, 1911) [H] 
Nearctic. Mixed coniferous woods; alder thickets. 
Little Grand Lake 27Jul94¢. 


Scotinotylus pallidus (Emerton, 1882) 

Nearctic. Sphagnum bogs; Empetrum barrens. 

Baie Verte Peninsula 220ct85° (CNC); Main River 16Aug85° 
2 (CNC); Pasadena 13, 19Jul84° 93 (CNC); Pinchgut Lake 
17Aug84? (CNC); Plumb (=Plum) Point 18Oct85° (CNC). 


Scotinotylus sacer (Crosby, 1929) 

Holarctic. Mixed coniferous woods; sphagnum bogs. 
Gander 28Jun98¢; Glide Lake 00Jul94°; Main River west 
OOJul98¢; Norris Point 27Jul982 , 27Jun-26Jul00e oc. 


Scyletria inflata Bishop & Crosby, 1938 
Nearctic. Mixed coniferous woods. 
Main River west 22Jul00¢; Burnt Cape 00Aug03°. 


Semljicola obtusus (Emerton, 1915) 
Nearctic. Mixed coniferous woods; Empetrum barrens. 
_ Main River west 22Jul00° ; Burnt Cape 00Aug03¢. 


Sisicottus montanus (Emerton, 1882) [H] 

Nearctic. Mixed coniferous woods. Common. 

Berry Head Pond (GMNP) 27Jul-16Aug99° 9°; Big Brook 
22-24Jun02° 9; Gander 1Jul78°; Lobster Cove 20Aug99 ° 
SO; Port au Choix 1Aug99¢. 


Sisicottus quoylei Miller, 1999 

Nearctic. Mixed coniferous woods. 

Logy Bay 11Jul02°; Long Pond (St. John’s) 10Jan99°<o; 
Main River west OOJul98¢. 


Sisicus apertus (Holm, 1939) 

Holarctic. Mixed coniferous woods. 
Bakers Brook estuary 8-27Aug00°°; 
25Jul92° 3; Glide Lake 00Jul94¢. 


Sisicus penifusifer Bishop & Crosby, 1938 
Nearctic. Sphagnum bogs. 
Rocky Harbour (Bottom Brook bog) 13Jul98°. 


Sisis rotundus (Emerton, 1925) 
Nearctic. Mixed coniferous woods. 
Glide Lake 00Jul94°; Main River west OOJul98°2c. 


Souessa spinifera (O. Pickard-Cambridge, 1874) [H] 
Nearctic. Mixed coniferous woods. 

Grandy Brook, Cow Head, Stanford River, Gambo: 21- 
30Jun49° ; 21-31 Jul49°; 1-31 Aug49° ° (FMNH?). Dates not 
connected to localities; year assumed 1949 from context (Hack- 
man 1954). Not found by CDD in Hackman’s collections in 
FMNH. This record may be erroneous but is retained here 
pending clarification. 


Corner Brook 


Sougambus bostoniensis (Emerton, 1882) [H; Hilaira algida 
Hackman, 1954] 

Nearctic. Mixed coniferous woods. 

Grand Bruit 13Jun49°° (FMNH and CNC); Lomond 
15Jul49° (FMNH),. 


Soulgas corticarius (Emerton, 1909) 
Nearctic. Mixed coniferous woods; litter at back of beaches. 
Bakers Brook estuary 27Aug98°. 


Stemonyphantes blauveltae Gertsch, 1951 

Nearctic. Mixed coniferous woods; Empetrum barrens. 
Cupids 5Sep99° ; Logy Bay 11Nov02°; Long Pond (St. John’s) 
12May80°; Mt. Scio O0OAug95°; St. John’s 21 Feb99¢. 


PICKAVANCE AND DONDALE: SPIDERS Of NEWFOUNDLAND 


265 


Styloctetor stativus (Simon, 1881) [H; Spirembolus orei- 
noides Chamberlin, 1948] 

Nearctic. Mixed coniferous woods. 

Kittys Brook 18Aug49?° (FMNH7?). 

Not found by CDD in Hackman’s collections in FMNH. This 
record may be erroneous but is retained here pending clari- 
fication. 


Tapinocyba bicarinata (Emerton, 1913) 
Nearctic. Mixed coniferous woods. 
Burnt Cape 00Aug03°; Main River west 00Jul98¢. 


Tapinocyba flagellata (Emerton, 1911) 

Nearctic. Mixed coniferous woods. 

Grandys Lake north 20Sep84¢° (CNC); Barachois Brook 
15Aug84? (CNC). 


Tapinocyba lindrothi Hackman, 1954 [H] 
Nearctic. Mixed coniferous woods. 
South Branch (Codroy) 3Jul49° (CNC). 


Tapinocyba minuta (Emerton, 1909) 

Nearctic. Alpine Empetrum barrens; sphagnum bogs. 

Burnt Cape 00Aug03° ° 9°; Killdevil Mountain 17Aug99°? 
3G; Point Riche 24Jul00°; Sally’s Cove 27Jul99¢. 


Tapinocyba simplex (Emerton, 1882) [H] 

Nearctic. Mixed coniferous woods. 

Bakers Brook estuary 8-27Aug00° 9; Glide Lake 00Jul94° 
3; Lobster Cove 21Aug98°; Stanford River 29Jul99° 9. 


Tapinopa bilineata Banks, 1893 

Nearctic. Litter at back of beach. 

Bakers Brook estuary 8-27Aug?; Blackhead (St. John’s) 
12Nov02° ; Green Point 26Aug98° . 


Tennesseellum formicum (Emerton, 1882) 
Nearctic. Mixed coniferous woods. 
Main River west 31 Jul00°; Norris Point 11Aug99°. 


Thyreosthenius parasiticus (Westring, 1851) [H] 
Holarctic. Mixed coniferous woods. 
Norris Point 28Jun-22Jul00°2 C. 


Tiso vagans (Blackwall, 1834) 
Palaearctic; introduced. Coastal Empetrum barrens. 
Blackhead (St. John’s) 31May02°¢. 


Tunagyna debilis (Banks, 1892) [H] 

Holarctic. Mixed coniferous woods; Empetrum barrens; shrubs 
at lower elevations of mountains. 

Big Brook 25Jun02° ° 99; Blackhead (St. John’s) 21 Apr99¢; 
Nameless Cove 16Aug01° °; Port au Choix1Aug99°; Table 
Point 23Jul97°. 


Typhocrestus pygmaeus (S¢rensen, 1898) 
Nearctic. Exposed coastal barrens. 
Burnt Cape OO0Jul03¢. 


Vermontia thoracica (Emerton, 1913) 
Holarctic. Mixed coniferous woods. 
Main River west OOJul98°2 Cc. 


Wabasso cacuminatus Millidge, 1984 
Holarctic. Mixed coniferous woods. 
Main River west OOJul98C. 


Wabasso quaestio (Chamberlin, 1949) 

Holarctic. Mixed coniferous woods; rocky coastal barrens. 
Big Brook 21Jun02°; Burnt Cape 00Aug03° 9; Main River 
west 00Jul98dc. 


266 


Walckenaria arctica Millidge, 1983 [H; Oedothorax montif- 
erus (Emerton, 1882)] 

Nearctic. Mixed coniferous woods. 

Bakers Brook estuary 8-27Aug00°°; Berry Head Pond 
(GMNP) 27Jul99¢; Glide Lake 00Jul94¢. 


Walckenaria atrotibialis (O. Pickard-Cambridge, 1878) [H; 
Entelecara abrupta (Emerton, 1909)] 

Holarctic. Mixed coniferous woods. 

Bakers Brook estuary 8-27Aug00°°; Berry Head Pond 
(GMNP) 28Jul99°; Norris Point 26Jul-14Aug00¢9; Port 
au Choix 1 Aug99°; Sally’s Cove 31Jul00¢. 


Walckenaeria auranticeps (Emerton, 1882) [H; Cornicular- 
ia auranticeps Emerton, 1882] 

Holarctic. Mixed coniferous woods. 

Gaff Topsail 20Aug49° (FMNH). 


Walckenaeria castanea (Emerton, 1882) [H; Trachynella 
nudipalpis (Westring, 1851), Minyriolus castaneus (Emerton)]| 
Nearctic. Mixed coniferous woods; coastal Empetrum barrens; 
sphagnum bogs. 

Berry Head Pond (GMNP) 28Jul-10Aug99° 9; Big Brook 
22Jun02° ° ; Rocky Harbour (Bottom Brook bog) 10Aug99° ; 
Point Riche 13Jul99° °; Port au Choix 8Jul00°S. 


Walckenaeria clavipalpis Millidge, 1983 

Nearctic. Mixed coniferous woods. 

Main River west OOJul989.°; Port au Choix 13Jul- 1Aug99°c, 
24Jul00¢; Stanford River 29Jul99° . 


Walckenaeria communis (Emerton, 1882) 

Nearctic. Coastal rocky barrens; sphagnum bogs; mixed conif- 
erous woods. 

Big Brook 20Jun02° ; Burnt Cape 00Aug03° 3; Eddies Cove 
East 21Jun00°; Point Riche 8-24Jul00° °; Sally’s Cove 
273ul99° . 


Walckenaeria cuspidata (Crosby & Bishop, 1931) [H; Cor- 
nicularia cuspidata Crosby & Bishop)] 

Nearctic. Rocky barrens; back of beaches. 

Big Brook 18Jun02° ° ; Nameless Cove 16Aug01°; Savage 
Cove 14Jul01°. 


Walckenaeria digitata (Emerton, 1913) 
Nearctic. Mixed coniferous woods. 
Main River west 22Jul00¢. 


Walckenaeria directa (O. Pickard-Cambridge, 1874) 
Nearctic. Mixed coniferous woods. 
Norris Point 13Jul00°. 


Walckenaeria exigua Millidge, 1983 [H; Cornicularia min- 
uta Emerton, 1882] 

Nearctic. Mixed coniferous woods; sphagnum bogs. 

Berry Head Pond (GMNP) 4Aug99¢; Rocky Harbour (Bot- 
tom Brook bog) 13Aug98° ; Gander 28Jul989°; Glide Lake 
OOJul942° 5; Point Riche 3Aug00°. 


Walckenaeria karpinskii (O. Pickard-Cambridge, 1873) [H; 
Cornicularia karpinskii (O. Pickard-Cambridge)] 

Holarctic. Mixed coniferous woods. 

Big Brook 16Jul01° °, 18Jul029; Burnt Cape O0Aug03° 3; 
Cape Raven 15Aug99¢; Main River west 00Jul98° 2. 


Walckenaeria lepida (Kulezynski, 1885) [H; Cornicularia 
unicornis (O. Pickard-Cambridge, 1861)] 

Holarctic. Mixed coniferous woods. 

Noel Pauls Brook 26Jun77° (CNC). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Walckenaeria pallida (Emerton, 1882) 
Nearctic. Mixed coniferous woods. 
Norris Point 27Jul00° . 


Walckenaeria palustris Millidge, 1983 
Nearctic. Mixed coniferous woods. 
Stanford River 29Jul99° . 


Walckenaeria redneri Millidge, 1983 
Nearctic. Mixed coniferous woods. 
Main River west OOJul98oc. 


Walckenaria spiralis (Emerton, 1882) [H; Cornicularia vig- 
ilax (Blackwall, 1853)] 

Holarctic. Sphagnum bogs; mixed coniferous woods. 
Bakers Brook estuary 3Jul98°; Rocky Harbour (Bottom 
Brook bog) 20Aug99° ; Burnt Cape O0OAug03°¢; Mt. Scio 
O0AuUg9S¢. 


Walckenaeria tricornis (Emerton, 1882) [H; Tigellinus tri- 
cornis (Emerton) 

Nearctic. Mixed coniferous woods. 

Burnt Cape O0Aug03°; Main River west 00Jul98¢; Stanford 
River 29Jul99¢c. 


Wubana drassoides (Emerton, 1882) 

Nearctic. Mixed coniferous woods. 

This record is from a single female. Identification of isolated 
females in this genus is difficult. Retained here as a New- 
foundland record pending confirmation by a male. 

Tuckers Head 30ct98°. 


Wubana pacifica (Banks, 1896) 

Nearctic. Mixed coniferous woods. 

A record based on both males and females. 

Big Brook 22Jun029; Burnt Cape O0Aug03° ; Corner Brook 
13Jul93°; Norris Point 29Jun-25Aug? co. 


Zornella cultrigera (L. Koch, 1879) [H]. 

Holarctic. Mixed coniferous woods. 

Nearctic records of Zornella are likely an undescribed species 
(Buckle et al. 2001), but the name is retained here pending 
revision of the genus. 

Main River west O00Jul98° . 


Linyphiidae CNC#76 

Not attributed to genus or species 

Mixed coniferous woods. 

Corner Brook 25Jul92° ; Glide Lake 00Jul94° ; Main River 
west 0O0Jul99° . 


LIOCRANIDAE (1 sp.) 

Agroeca ornata Banks, 1892. 

Holarctic. Mixed coniferous woods. 

Big Brook 21Jun02°; Burnt Cape O0Aug03°°; Gander 
28Jul98° ; Glide Lake OOJul96° ° ; Main River west OOJul98° . 


LYCOSIDAE (24 spp.) 

Alopecosa aculeata (Clerck, 1758) [H] 

Holarctic. Rocky and Empetrum barrens; sphagnum bogs. 
Brunette Island 22Jun80¢; Burnt Cape 00Aug03°°; Cape 
Race 11Jun95°; Hawke Hills 2Jun95°°9°; Point Riche 
4Aug00°2 °. 


Arctosa alpigena (Doleschall, 1852) [H] 

Holarctic. Exposed coastal Empetrum barrens; tops of the 
Long Range Mountains. 

Big Brook 22Jun02°9; Burnt Cape 00Aug03° 29°; Gros 
Morne Mountain 17Aug94° ; Killdevil Mountain 17Aug99° ° ; 
Watts Point 9Aug95d¢. 


2005 


Arctosa insignita (Thorell, 1872) 
Holarctic. Mixed coniferous woods. 
Main River west 00Aug98°. 


Arctosa lama Dondale & Redner, 1983 [H; Arctosa virgo 
(Chamberlin, 1925)] 

Nearctic. Sphagnum bogs. 

Grandy Brook 25Jun49° (FMNH). 


Arctosa raptor (Kulczyfski, 1885) [H; Arctosa quinaria 
(Emerton, 1895)] 

Holarctic. Mixed coniferous woods; sphagnum bogs. 

Glide Lake 19Aug84° (CNC); North Arm Mountain (Bonne 
Bay) 13Aug87° °? (CNC). Cow Head, Grand Bank, Lomond: 
11-20Jun49c; 1-31Jul49°°cqc; 1-31Aug?°S (FMNH). 
Dates not connected to localities; year assumed 1949 from 
context (Hackman 1954). 


Arctosa rubicunda (Keyserling, 1877) [H; Arctosa emertoni 
Gertsch, 1934] 

Nearctic. Sphagnum bogs. 

Cape Ray 1Jul96° ? (and CNC). 


Pardosa albomaculata Emerton, 1885 

Nearctic. Talus and mixed vegetation at higher elevations of 
the Long Range Mountains. 

Gros Morne Mountain 8Oct94° ; Tablelands 15Aug92°. 


Pardosa concinna (Thorell, 1877) [H] 

Nearctic. Exposed coastal Empetrum barrens. Common. 
Cape Onion 4Aug96° °; Cape Ray 1Jul96¢; Cape Spear 3Sep 
95°C; Point Riche 17Aug98° °3S; Watts Point 1Aug95°c. 


Pardosa furcifera (Thorell, 1875) [H] 

Nearctic. Empetrum barrens on exposed coasts and at high- 
er elevations. 

Big Brook 21Jun02°°; Killdevil Mountain 17Aug99°<c; 
Nameless Cove 16Aug01°; Savage Cove 14Jul01°; Watts 
Point 19Aug95° °co, 


Pardosa fuscula (Thorell, 1875) [H] 

Nearctic. Sphagnum bogs. 

Bay Bulls 15Aug96°°; Big Brook 21Jun02°; Burnt Cape 
OO0Aug03° ° So; Cape Freels 12Jul01; Cape Ray 1Jul96°. 


Pardosa groenlandica (Thorell, 1872) [H] 

Nearctic. Cobble and pebble beaches. In NF rarely found away 
from the coast. Common. 

Bay Bulls 1May96°; Bristols Hope 11Nov94°; Conche 
16Jul03°; Port au Port Peninsula? o; St. Pauls 6Aug95°. 


Pardosa hyperborea (Thorell, 1872) [H; Pardosa saltuaria 
(L. Koch, 1870)] 

Holarctic. Mixed coniferous woods; Empetrum barrens; sphag- 
num bogs. Common. 

Boutitou 17Jul039; Cape Ray 1Jul96° °C; Cape Spear 
3Sep95°S9; Hawke Hills 2Jun95°°; Killdevil Mountain 
17Aug992 ° , 


| Pardosa lapidicina Emerton, 1885 [H] 

| Nearctic. Talus slopes of the Long Range Mountains. 

| Gros Morne Mountain 31Jul95° ; Killdevil Mountain 2Jul95°9 ; 
| Tablelands 11 Aug94°. 


| Pardosa mackenziana (Keyserling, 1877) [H] 

| Nearctic. Mixed coniferous woods. 

| Bakers Brook 8-27Aug00° 9°; Gander 1Jul79¢; Glide Lake 
| 8-22Jul?; Port au Choix13Jul99° ; Woody Point 90ct94°. 


PICKAVANCE AND DONDALE: SPIDERS Of NEWFOUNDLAND 


267 


Pardosa moesta Banks, 1892 [H] 

Nearctic. Coastal and inland Empetrum barrens. 

Cape Spear 3Sep95° °; Gander 21Jun98¢9°; Green Gardens 
18Jun94°°C0; Hawke Hills 2Jun95°°; Port au Choix 
6Aug9I62 °. 


Pardosa xerampelina (Keyserling, 1877) [H] 

Nearctic. Mixed coniferous woods; Empetrum barrens; sphag- 
num bogs. 

Glide Lake 7Jul94° 0; Green Gardens 7Jul97° ; Port au Choix 
23Jul97¢; Tablelands 23Jul93° 9; TNNP 11Jul93°. 


Pirata bryantae Kurata, 1944 [H] 
Nearctic. Mixed coniferous woods. 
Main River west O0Aug98°¢. 


Pirata canadensis Dondale & Redner, 1981 

Nearctic. Mixed coniferous woods; sphagnum bogs. 

Rocky Harbour (Bottom Brook bog) 29Jul98°, 20Aug99¢ ; 
Main River west O0Jul98°¢. 


Pirata cantralli Wallace & Exline, 1978 [H; Pirata insularis 
Emerton, 1885] 

Nearctic. Sphagnum bogs. 

Cape Broyle, Gander, Grand Bruit, Pushthrough, Woody Point: 
1-30Jun49? 9d; 1-31Jul49 °°; 1-31Aug49°° (FMNH). 
Dates not connected to localities; year assumed 1949 from 
context (Hackman 1954). 


Pirata insularis Emerton, 1885 [H] 

Holarctic. Sphagnum bogs. 

Bakers Brook estuary 9Jul99°; Berry Head Pond (GMNP) 
22Aug00° 9; Rocky Harbour (Bottom Brook bog) 23Jul99° 3; 
Mt. Scio O0Aug95¢. 


Pirata minutus Emerton, 1885 [H] 
Nearctic. Mixed coniferous woods; sphagnum bogs. 
Main River west 18Jun99°. 


Pirata montanus Emerton, 1885 [H] 

Holarctic. Mixed coniferous woods; sphagnum bogs. 

Base of Killdevil Mountain 13Jul49° (FMNH); East Brook 
(which one unspecified) 14Jul49° ° (FMNH). 


Pirata piraticus (Clerck, 1758) [H] 

Holarctic. Sphagnum bogs. 

Bay Bulls 5Aug96°; Big Brook 24Jun02° 9°99; Cape Ray 
1Jul96°; Hawkes Bay 1Jul99°. 


Trochosa terricola Thorell, 1856 |[H: Trochosa terricola 
pratensis Emerton, 1885] 

Holarctic. Mixed coniferous woods; shrubs and herbs in clear- 
ings of woods; litter and stones at back of beaches; abandoned 
meadows; gardens. Common. 

Brunette Island 22Jun80°; Nameless Cove 16Aug01°; Nor- 
ris Point 9Aug95°; Shallow Bay 27May96°; St. John’s 
12May80° °. 


MIMETIDAE (1 sp.) 

Ero canionis Chamberlin & Ivie, 1935 [H] 

Nearctic. Empetrum barrens; shrubs and herbs at edges of 
woods. 

Point Riche 17Jul00°. 


NESTICIDAE (1 sp.) 

Eidmanella pallida (Emerton, 1875) [H; Nesticus pallida 
Emerton] 

Palaearctic; introduced. Mixed coniferous woods; caves: 
human habitations. 

Corner Brook (in cave) 13Sep88° (CNC). 


268 


PHILODROMIDAE (7 spp.) 

Philodromus alascensis Keyserling, 1884 [H; Philodromus 
pernix Blackwall, 1846] 

Holarctic. Exposed coastal Empetrum barrens. 

Boutitou 17Jul03°. 


Philodromus imbecillus Keyserling, 1880. 

Nearctic. Mixed coniferous woods. 

Bay d’Espoir 11Jul88°¢o° (CNC); Witch Hazel Ridge 
25Jun81° ° (CNC). 


Philodromus mysticus Dondale & Redner, 1975 
Holarctic. Mixed coniferous woods. 
Main River west OOJul98°. 


Philodromus placidus Banks, 1892 
Nearctic. Mixed coniferous woods. 
Gander 00Aug98o°. 


Philodromus rufus quartus Dondale & Redner, 1968 [H; 
Philodromus rufus Walckenaer, 1826] 

Nearctic. Mixed coniferous woods; shrubs and herbs in clear- 
ings in woods. 

Gadds Harbour 24Jul99°; Gander 1Jul78° °°; Logy Bay 
8-23Jul01°S; Norris Point 3Jul00°; Rocky Harbour 30Jul97°. 


Thanatus formicinus (Clerck, 1758) [H] 

Holarctic. Exposed coastal rocky and Empetrum barrens. 
Bay Bulls 4May96°; Burnt Cape 00Aug03°; Logy Bay 
12Jun01°; Table Point 3Aug95°; Watts Point 9Aug95°. 


Tibellus maritimus (Menge, 1875) [H] 

Holarctic. Coastal Empetrum barrens. 

Bay Bulls 16Jul95¢; Logy Bay 8Jul01°; Norris Point 3Jul00¢; 
St. John’s 12May80° ; St. Lunaire O0Aug00°. 


PHOLCIDAE (1 sp.) 

Pholcus phalangioides (Fuesslin, 1775) 
Palaearctic; introduced. Houses in St. John’s. 
St. John’s 15Aug00°. 


PISAURIDAE (1 sp.) 

Dolomedes striatus Giebel, 1869 [H: Dolomedes fulviatrona- 
tus Bishop, 1924 and Dolomedes vittatus Walckenaer, 1837] 
Nearctic. Edges of small pools; wet areas. _ 

Come by Chance 27Aug49? (FMNH); Corner Brook 16Aug 
49° (FMNH); East Brook, Lomond 15Jul49° (FMNH); 
Hogans Pond 6Jun49° (FMNH). 


SALTICIDAE (15 spp.) 

Chalcoscirtus alpicola (L. Koch, 1876) 

Holarctic. Sub-arctic, exposed, coastal rocky and Empetrum 
barrens. 

Big Brook16Jul01¢. 


Dendryphantes nigromaculatus (Keyserling, 1885) 
Nearctic. Exposed coastal barrens; exposed stunted woods. 
Bay d’Espoir 15Jul85° ° (CNC). 


Evarcha hoyi (Peckham & Peckham, 1883) [H] 

Nearctic. Empetrum barrens; talus slopes. 

Boutitou 17Jul039; Gander O0Aug98°; Killdevil Mountain 
9Aug94c; Logy Bay 16Sep03°; Main River west 29Jul00¢. 


Ghelna canadensis (Banks, 1897) 
Nearctic. Sphagnum bogs. 
Bakers Brook estuary 19Aug99°. 


Neon nellii Peckham & Peckham, 1888 [H] 
Nearctic. Rocky and Empetrum barrens. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Berry Head Pond (GMNP) 19Jul00¢; Logy Bay 18Jun019; 
Norris Point 28Jul00¢; Point Riche 17Jul002 °. 


Pelegrina flavipes (Peckham & Peckham, 1888) [H; Meta- 
phidippus flavipedes (Peckham & Peckham)] 

Nearctic. Mixed coniferous woods; shrubs and herbs in clear- 
ings in woods. 

Cupids 5Sep99° 9; St. John’s 8Sep98¢. 


Pelegrina montana (Emerton, 1891) [H; Metaphidippus mon- 
tanus (Emerton)| 

Nearctic. Empetrum barrens; low vegetation. 

Brunette Island 14Jul94°; Gander 1Jul78°°; Millertown 
26Jun80° °; St. John’s 8Sep98°. 


Phidippus borealis Banks, 1895 [H; Phidippus purpuratus 
Keyserling, 1885] 

Nearctic. Talus slopes and rocky barrens at low and medium 
elevations in the Long Range Mountains. 

Tablelands 11Aug94°; Winter House Brook canyon 10Jul97¢. 


Phidippus whitmanii Peckham & Peckham, 1909 [H; Phi- 
dippus princeps (Peckham & Peckham, 1883] 

Nearctic. Empetrum barrens; sphagnum bogs. 

Rencontre West 15Jun49° (FMNH). 


Salticus scenicus (Clerck, 1758) [H] 

Palaearctic; introduced. On houses; Empetrum barrens; aban- 
doned meadows; litter and stones at back of beaches. 
Embree 12Nov 92°; Clarkes Beach 24May98¢; Logy Bay 
18Jun019°; Norris Point 29Jul00¢; St. John’s 2Jun93c. 


Sitticus cutleri Proszynski, 1980 
Nearctic. Coastal Empetrum barrens. 
Maddox Cove 1Jul94¢. 


Sitticus finschii (L. Koch, 1879) 
Holarctic. Mixed coniferous woods; sphagnum bogs. 
Gander 00Aug02°; Main River west 8Aug00°. 


Sitticus floricola palustris (Peckham & Peckham, 1883) [H; 
Sitticus palustris (Peckham & Peckham)] 

Nearctic. Back of beaches; sand dunes; edge of lakes among 
driftwood. 

Broom Point 16Jul98° 3; Conche 27Jul04° 3; Gadds Harbour 
8Jul9400; Outer Cove 3Aug969; St. Pauls 25Jul97° °. 


Sitticus ranieri (Peckham & Peckham, 1909) 

Holarctic. Empetrum barrens; talus slopes. 

Gander 00Aug98° SS; Logy Bay 12Jun01°°; Killdevil Moun- 
tain 17Aug99¢c. 


Sitticus striatus Emerton, 1911 [H] 

Holarctic. Mixed coniferous woods; sphagnum bogs. 
Blackhead (St. John’s) 10Jun01°; Lomond 14Jul98° ; Main 
River west 27Jul00° °c. 


TETRAGNATHIDAE (6 spp.) 

Meta ovalis (Gertsch 1933) [H; Meta menardi (Latreille, 
1804)] 

Nearctic. Houses and outbuildings; basements; root cellars. 
Colliers 14Dec04° ; Logy Bay 0OOMay98°; St. John’s OOJun 
on. 


Pachygnatha brevis Keyserling, 1884 [H] 
Nearctic. Shrubs and herbs in damp places; near waterways. 
St. John’s 12may809¢. 


Tetragnatha elongata Walckenaer, 1842 [H] 
Nearctic. Shrubs and herbs in damp places; near waterways. 
Portugal Cove 11Jul79°. 


2005 PICKAVANCE AND DONDALE 


Tetragnatha extensa (Linnaeus, 1758) [H] 

Holarctic. Shrubs and herbs in damp places; near waterways; 
wetlands; herbs at back of beaches; road-cuttings. 

Hawkes Bay 1Jul99°°¢; Plum Point 3Jul80°; Port au Choix 
4Aug97°S; Raleigh 4Aug98° 9; Rocky Harbour 30Jul97°. 


Tetragnatha laboriosa Hentz, 1850 [H] 

Nearctic. Shrubs and herbs in damp places; near waterways. 
Rocky Harbour (Bottom Brook bog) 16Jul00¢; Gander 
28Jul98. 


Tetragnatha versicolor Walckenaer, 1842 

Nearctic. Shrubs and herbs in damp places; near waterways. 
Bakers Brook estuary 8-27Aug00°°; Berry Head Pond 
(GMNP) 24Jul99¢. 


THERIDIIDAE (26 spp.) 

Achaearanea ohlerti (Thorell, 1870) 
Holarctic. Mixed coniferous woods. 
Gadds Harbour 24Jul99° . 


Achaearanea tabulata Levi, 1980 

Introduced; of unknown origin (Dondale et al. 1994). 

In and on houses; on sheds and outbuildings; in gardens. 
St. John’s 24Sep99°, 3AugO01°, 23Aug01°. 


Achaearanea tepidariorum (C. L. Koch, 1841) 

Introduced, of unknown origin; possibly tropical (Lindroth 
1957). 

In and on houses; on sheds and outbuildings; in gardens. 

St. John’s 3Aug01¢, IMay04° °. 


Crustulina sticta (O. Pickard-Cambridge, 1861) [H; Crus- 
tulina borealis Banks, 1900] 

Holarctic. Among stones at back of beaches; among stones 
on Empetrum barrens; among stones on sides and tops of 
mountains. 

Eddies Cove East 21Jun01° °°; Killdevil Mountain 17Aug 
99°°; New Ferolle 16Jul00° °; Point Riche 29Jun99°<c; 
Port au Choix5Jul04° °c, 


Dipoena nigra (Emerton, 1882) 
Nearctic. Mixed coniferous woods; shrubs. 
Gander 23Jun82¢ (CNC). 


Enoplognatha caricis (Fickert, 1876) [H; Theridion tectum 
(Keyserling, 1884)] 

Holarctic. Mixed coniferous woods; shrubs; rocky barrens; 
coastal Empetrum barrens. 

Point Riche 17Jul00°°; Port au Choix 4Jul04° ° (and 
CNC); Table Point 23Jul97° ° (and CNC). 


Enoplognatha intrepida (S¢rensen, 1898) 

Nearctic. Exposed rocky and Empetrum barrens. 

Burnt Cape 00Aug039°; Cape Raven 15Aug99°<C; Point 
Riche 24Jul98°c. 


Enoplognatha marmorata (Hentz, 1850) 

Nearctic. Mixed coniferous woods; sand dunes. 

Cape Freels 10Jul019; Glide Lake 00Jul96°; Table Point 
18-30Jul00c. 


Enoplognatha ovata (Clerck, 1758) 

| Palaearctic; introduced. Shrubs and herbs; roadside vegeta- 
tion; gardens; coastal barrens. 

Gadds Harbour 24Jul99° °; Rocky Harbour 8Aug97°; St. 
John’s 27Aug97° °; Woody Point 8Aug93°. 


Neottiura bimaculata (Linnaeus, 1767) 

Palaearctic; introduced. Mixed coniferous woods; gardens. 
Gadds Harbour 24Jul99°; Gander O0Jul98¢; Logy Bay 
24Jul03¢c. 


: SPIDERS of NEWFOUNDLAND 


269 


Robertus banksi (Kaston, 1946) [H; Ctenium banksi Kaston] 
Nearctic. Mixed coniferous woods. 

Cow Head, Deer lake, Lomond, South Branch (Codroy), 
Stephenville Crossing: 1-20Jun49°? 9°90; 1-31Jul49° °; 11- 
20Aug49? (FMNH). Dates not connected to localities; year 
assumed 1949 from context (Hackman 1954). 


Robertus borealis (Kaston, 1946) [H; Ctenium boreale Kaston] 
Nearctic. Mixed coniferous woods. 

Gaff Topsail 20Aug49° (FMNH); Cow Head 12Aug49°? 
(FMNH); 40 miles west of St John’s 6Aug76° (CNC). 


Robertus fuscus (Emerton, 1894) [H; Ctenium fuscum (Emer- 
ton)]} 

Nearctic. Empetrum barrens; rocky barrens. 

Boutitou 17Jul03°; Main River west OOJul98¢; Nameless 
Cove 16Aug01° °°; Point Riche 17Aug98°; Table Point 
30Sep98°. 


Robertus riparius (Keyserling, 1886) [H; Ctenium riparium 
(Keyserling)] 

Nearctic. Mixed coniferous woods. 

Cing Cerf Brook, Grand Bruit, Grandy Brook, South Branch 
(Codroy), Table Mountain (St. Georges): 11-20, 21-30Jun 
492200 (FMNH); 1-10, 11-20, 21- 31Jul49° 9° (FMNH); 
11-20Aug49° (FMNH). Dates not connected to localities; 
year assumed 1949 from context (Hackman 1954). 


Rugathodes aurantius (Emerton, 1915) [H; Theridion auran- 
tium Emerton] 

Holarctic. Mixed coniferous woods. 

Corner Brook 19Jul93°; Port au Choix 24Jun00¢. 


Rugathodes sexpunctatus (Emerton, 1882) [H; Theridion 
sexpunctatum Emerton] 

Holarctic. Mixed coniferous woods; shrubs and herbs; gardens. 
Berry Head Pond (GMNP) 4Aug99° 2°; Conche 16Jul03° ° ; 
base of Killdevil Mountain 20Jul00°; Norris Point 13Aug 
94°; Port au Choix 24Jul00¢. 


Steatoda bipunctata (Linnaeus, 1758) [H] 

Palaearctic; introduced. On or near human structures: houses, 
basements, outbuildings and fences. 

Corner Brook 8Jul99°; Deer Lake 12Sep97°; Dunville 
25Apr97¢; Port au Choix 4Jul03¢; St. John’s 10Jan02°. 


Theonoe stridula Crosby, 1906 

Nearctic. Mixed coniferous woods; wetlands. 

Berry Head Pond (GMNP) 19Jul00° 0; Rocky Harbour (Bot- 
tom Brook bog) 6Aug98°; Main River west 22Jul00¢. 


Theridion differens Emerton, 1882 

Nearctic. Mixed coniferous woods; wetlands. 

Gander 20Jul78° (CNC); South Branch (Codroy) 24- 31Jul 
745 (CNC); St. Georges 1Jul87° (CNC). 


Theridion glaucescens Becker, 1879 [H] 

Nearctic. Mixed coniferous woods; wetlands. 

Woody Point 17Jul49° (FMNH?); Gambo 25Aug49° 
(FMNH?). This species not found by CDD in Hackman’s 
collections in FMNH. This record may be erroneous but is 
retained here pending clarification. 


Theridion montanum Emerton, 1882 [H] 

Holarctic. Mixed coniferous woods. 

Berry Head Pond (GMNP) 24Jul99¢; Gadds Harbour 
24Jul99° ; Mt. Scio 15Sep99° ; Shallow Bay 13Jul97¢. 


Theridion murarium Emerton, 1882 [H] 

Nearctic. Mixed coniferous woods; wetlands. 

Square Pond (which one unspecified) 9Jun81¢ (CNC). All 
Hackman’s specimens immature. 


270 


Theridion petraeum L. Koch, 1872 

Holarctic. Among stones at mid and high elevations on the 
Long Range Mountains. 

Tablelands 1 Aug0Q0° °. 


Theridion pictum (Walckenaer, 1802) 
Holarctic. Mixed coniferous woods. 
Gander 00Aug98°; Main River west 28Jul00¢. 


Theridula emertoni Levi, 1954 [H; Theridula sphaerula Emer- 
ton, 1882] 

Nearctic. Mixed coniferous woods. 

Gander 00Aug98°; Lomond 14Jul98°. 


Thymoites oleatus (L. Koch, 1879) - 
Holarctic. Exposed coastal Empetrum barrens. 
Savage Cove 23Aug98°. 


THERIDIOSOMATIDAE (1 sp.) 

Theridiosoma gemmosum (L. Koch, 1877) [H; Theridioso- 
ma radiosum (Emerton, 1884)] 

Palaearctic; introduced. Mixed coniferous woods; wetlands. 
Bay d’Espoir 15Jul85° (CNC); South Branch (Codroy) 
3Jul49° ° (FMNH); Spruce Brook 8-9Jul49° (FMNH). 


THOMISIDAE (14 spp.) 

Bassaniana utahensis (Gertsch, 1932). [H; Coriarachne 
versicolor Keyserling, 1880] 

Nearctic. Mixed coniferous woods; shrubs and herbs in 
clearings. 

Gander 1Jul78°; Terra Nova village 15Sep00°. 


Misumena vatia (Clerck, 1758) [H] 

Holarctic. On flowers among shrubs and herbs; on flowers 
along roadsides. 

Badger 27Jun80° ; Bakers Brook estuary 26Aug98° ; Black- 
head (St. John’s) 18Sep03°; Gander 1Jul79°; Hawkes Bay 
1Jul9sc. 


Ozyptila distans Dondale & Redner, 1975 [H; Oxyptila 
americana Banks, 1895] 

Nearctic. Mixed coniferous woods. 

Gander 00Aug98°; TNNP 1 1Jun93°°¢. 


Ozyptila gertschi Kurata, 1944 
Holarctic. Coastal Empetrum barrens. 
Burnt Cape 00Aug03° °c. 


Ozyptila sincera canadensis Dondale & Redner, 1975 [H; 
Ozyptila bryantae Gertsch, 1939] 

Nearctic. Coastal rocky barrens. 

Burnt Cape 00Aug03° 2°99; Point Riche 13Jul99°°; Port 
au Choix 5Jul04°. 


Xysticus canadensis Gertsch, 1934 [H] 

Holarctic. Mixed coniferous woods. 

Crooked Lake 19Aug84? (CNC); Hampden 14Jun77? (CNC); 
Kings Point 19Aug84° (CNC); Main River 16Aug84° (CNC); 
20 km north of Millertown 19Aug849 (CNC). 


Xysticus discursans Keyserling, 1880 [H] 

Nearctic. Mixed coniferous woods. 

Gander 1Jul78°; Lomond 9Jun94°; Main River west OOJul 
989; Port au Choix 24Jul00¢. 


Xysticus elegans Keyserling, 1880 [H] 

Nearctic. Mixed coniferous woods; shrubs and herbs. 
Corner Brook 16Aug49° (FMNH); Gambo 28Aug49° 
(FMNH). 


Xysticus emertoni Keyserling, 1880 [H] 
Holarctic. Rocky barrens; back of beaches. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Green Gardens 27Jun95° ; Highlands of St. John 14Aug00°; 
New Ferolle 13Jul04¢; Point Riche 12Jul99° ; Western Brook 
estuary 29Jul97°? °. 


Xysticus keyserlingi Bryant, 1930 

Nearctic. Exposed coastal Empetrum barrens; tops of Long 
Range Mountains. 

Burnt Cape 00Aug03°°; Killdevil Mountain 17Aug99¢;; 
Point Riche 4Aug00°. 


Xysticus labradorensis Keyserling, 1887 
Holarctic. Exposed coastal Empetrum barrens. 
Point Riche 18Aug02°°9. 


Xysticus luctuosus (Blackwall, 1836) 

Holarctic. Mixed coniferous woods; shrubs and herbs. 
Barbace Cove 7Jul03¢; Main River west OOJul98¢; Seal Cove 
(Bonne Bay) 14Jul98°9. 


Xysticus obscurus Collett, 1877 
Holarctic. Exposed, rocky, coastal barrens; back of beaches. 
Big Brook 19-21Jun02°°. 


Xysticus triguttatus Keyserling, 1880 [H] 

Nearctic. Empetrum barrens; abandoned meadows. 

Big Brook 25Jun02¢; Blackhead (St. John’s) 9Jul01¢; Eddies 
Cove East 21Jun00°°; Logy Bay 11Jul02°°; Point Riche 
8Jul00c. 


ULOBORIDAE (1 sp.) 

Hyptiotes gertschi Chamberlin & Ivie, 1935 

Nearctic. Mixed coniferous woods. 

Deer Arm 8Aug97°; Berry Head Pond (GMNP) 14Aug98¢. 


Discussion 
Origins of the Newfoundland spider fauna 

For present purposes we call all unidentified New- 
foundland species Nearctic because the taxonomy of 
European and Holarctic spiders is relatively well known. 
The majority (c. 61%) of Newfoundland species are 
Nearctic, with significant Holarctic (c. 33%) and in- 
troduced (c. 6%) components (Table 1). Given that re- 
cruitment to Newfoundland would largely have been — 
from the west on the prevailing winds (see above), 
the Nearctic proportion is not surprising. In Canada in 
general, the proportion of Holarctic species appears 
to increase with increasing latitude. The wetlands of 
Quebec, for example, had an overall Holarctic propor- | 
tion of about 33%, while in the northern wetlands of | 
the subarctic and arctic parts of northern Quebec the 
Holarctic component was nearly 50% (Koponen | 
1994). On the Belcher Islands (Hudson Bay) the Hol- | 
arctic proportion was 58% (Koponen 1992). 


Introduced species 

The number and proportion of introduced Palae- 
arctic species from several different arthropod taxa | 
reflect Newfoundland’s long history of European im- — 
migration and commerce, but the percentage varies 
between taxa. For example, the Plecoptera (stoneflies) 
and Ephemeroptera (mayflies) have no introduced spe- 
cies (Larson and Colbo 1983), approximately 17% (20 
of 120) of the Cicadellidae (leafhoppers) are introduced 
(Hamilton and Langor 1987), and all 12 terrestrial © 
Isopoda (wood-lice) were introduced to Newfound- 
land (Palmen 1951). 


2005 


PICKAVANCE AND DONDALE: SPIDERS of NEWFOUNDLAND 


271 


TABLE |. Origins and composition of the spider fauna of Newfoundland. 


Holarctic 


0 
0 
10 


Family Nearctic 


Agelenidae 
Amaurobiidae 
Araneidae 
Clubionidae 
Cybaeidae 
Dictynidae 
Gnaphosidae 
Hahniidae 
Linyphiidae 
Liocranidae 
Lycosidae 
Mimetidae 
Nesticidae 
Philodromidae 
Pholcidae 
Pisauridae 
Salticidae 
Tetragnathidae 
Theridiidae 
Theridiosomatidae 
Thomisidae 
Uloboridae 


— 
mem OO 


ies) 


HSB ISON NOK OWOKF NON NAW DA — O~AITN 


— 


Nn 
OonNoworshypoorso dc OF WS 


Totals: 22 Families 223 (61.4%) 119 (2.8%) 


TABLE 2. The 21 spider species introduced to Newfoundland. 


Agelenidae 
Tegenaria atrica 
Tegenaria domestica 
Tegenaria duellica 
Araneidae 
Araneus diadematus 
Larinioides sclopetarius 
Zygiella atrica 


Linyphiidae 


Tiso vagans 


Pholcidae 


Pholcus phalangioides 


Twenty-one spider species are here considered to be 
introductions to Newfoundland (Table 2). Two species 
are widespread and locally abundant throughout the 
island: Araneus diadematus and Tegenaria domestica 
occur from the Avalon Peninsula in the south to the 
tip of the Northern Peninsula. Interestingly, neither 
of these now widespread species was seen by Hack- 
man (1954) who only included A. diadematus in his 
list on the basis of a mention in Wiehle (1931). In 
fact the first North American record for this species 
was from Brigus, Newfoundland (Pickard-Cambridge 
1881). Since both species are large and not easily over- 
looked, both were probably very rare prior to 1954 
and have become widespread in the last 50 years. A 
similar, rapid colonisation has been demonstrated for 
other arthropod taxa. For example, in 1955 the cara- 
bid beetles Amara aulica Panzer and Pterostichus 
melanarius Ulliger were respectively confined to the 


Centromerita bicolor 
Dicymbium nigrum 
Erigone dentipalpis 
Lepthyphantes leprosus 
Lepthyphantes tenuis 


Eidmanella pallida 


Introduced Total species % of total 
3 4 1.1 
0 5 1.4 
3 20 55 
0 14 3.9 
0 l <| 
0 9 25 
0 14 3.9 
) 6 1.7 
6 19] 52.6 
0 1 <l 
0 24 6.6 
0 | <] 
| | <I 
0 7 1.9 
] ] <l 
0 l <i 
1 15 4.1 
0 6 iy 
5 26 fi des | 
| 1 =a | 
0 14 3.9 
0 l <l 
21 (5.8%) 363 

Salticidae 
Salticus scenicus 
Theridiidae 


Achaearanea tabulata 
Achaearanea tepidariorum 
Enoplognatha ovata 
Neottiura bimaculata 
Steatoda bipunctata 
Theridiosomatidae 
Theridiosoma gemmosum 


west coast of Newfoundland and known only from a 
single known locality on the Avalon Peninsula. In less 
than 30 years both species became widespread across 
Newfoundland (Larson and Langor 1982). 

Other species are well established in Newfound- 
land but less widespread. Steatoda bipunctata has not 
been found north of Port au Choix. Neither Larin- 
loides sclopetarius nor Salticus scenicus have been 
found north of Rocky Harbour. Zygiella atrica is par- 
ticularly frequent on the Avalon Peninsula but has been 
found nowhere else in Newfoundland except Deer 
Lake. Whether these distributions are artefacts of col- 
lecting, intermediate points in the spread of these spe- 
cies across the island, or the limits of these species’ 
colonising ability is unknown. 

Dicymbium nigrum is well established as several 
wild populations but only in a restricted area in and 
around St. John’s. A few species are known only from 


272 THE CANADIAN FIELD-NATURALIST Vol. 119 
TABLE 3. Comparison of seven Canadian provincial and territorial spider faunas. 

BC! OC SK? AB? MB* NF YKS 
Area 1000 km2 ° 947.8 1700 651.9 661.19 649.95 111.39 483.45 
Ecodistricts’ 1i2 87 154 146 94 25 78 
Latitude range°N ° 49-60 45-62 49-60 49-60 49-60 47-52 60-69 
Total species 657 617 550 527 483 363 297 
Total families 35 28 23 22 20 22 IS 
Agelenidae S 4 9 8 8 “ 1 
Amaurobiidae 10 14 4 5 7 5 2 
Antrodiaetidae 2 0 0 0 0 0 0 
Anyphaenidae 2 0 1 l 0 0 0 
Araneidae 33, 31 26 29 27 20 16 
Clubionidae 15 21 17 19 20 14 (i! 
Corinnidae 5 3 3 2 0 0 0 
Cybaeidae el 1 0 0 l 0 
Dictynidae 29 pe) 30 19 23 9 10 
Dipluridae 0 0 0 0 0 0 
Dysderidae ] 1 0 0 0 0 0 
Gnaphosidae 49 34 46 52 31 14 23 
Hahniidae 15 5 6 | 5 6 4 
Linyphiidae 231 240 198 201 179 19] 136 
Liocranidae 5 8 2) 5 6 1 1 
Lycosidae 47 53 41 46 40 24 30 
Mecicobothriidae l 0 0 0 0 0 0 
Mimetidae 2 5 | l 2 1 0 
Miturgidae | 0 0 0 0 0 
Mysmenidae 1 0 0 0 0 0 0 
Nesticidae l 2 0 0 0 l 0 
Oecobiidae 0 l 0 0 0 0 0 
Oxyopidae 1 l | 0 0 0 
Philodromidae 33 20 24 22 22 7 13 
Pholcidae 3 | l l 0 0 
Pimoidae 2 0 0 0 0 0 0 
Pisauridae | 5 2 2 4 l l 
Salticidae 45 43 50 35 33 15 17 
Segestriidae l 0 0 0 0 0 0 
Telemidae l 0 0 0 0 0) 0 
Tetragnathidae 12 15 12 8 13 6 3 
Theridiidae 51 48 38 27 31 26 14 
Theridiosomatidae 0 l 0 0 0 | 0 
Thomisidae By, 32 52 34 29 14 1 
Titanoecidae 2 | 2 2 2 0 Z 
Uloboridae | 3 l | l l 0 
Zoridae | 0 0 0 0 0 0 


‘Bennett 2001; Bennett et al. 2004*. 7Bélanger and Hutchinson 1992, as revised by Paquin et al. 2001, with additions in 
Paquin and Dupérré 2003. *Buckle and Holmberg 2004*. *Aitchison-Benell and Dondale 1990. *Dondale et al. 1997. Some 
familial placements adjusted in accordance with Platnick 2005*. 

Government Websites: Alberta, British Columbia, Manitoba, Newfoundland and Labrador, Quebec, Saskatchewan, Yukon 


2004* (Some numbers approximate) 
’ Ecological Stratification Working Group 1995. 


one or two specific buildings in the St. John’s area, 
either houses and other domestic structures (Pholcus 
phalangioides, Tegenaria duellica, Achaearanea tep- 
idariorum and A. tabulata) or commercial buildings 
(Tegenaria atrica). The remaining species are known 
from small numbers of specimens from a small num- 
ber of localities in the wild; little can be said about 


them until further collections tell us more about their 
distributions and habitats. 


Composition of the fauna: comparisons with other areas 

In general terms the composition of any terrestrial 
fauna will be a reflection of land area, ecological diver- 
sity and latitudinal position. Table 3 compares these 


2005 


PICKAVANCE AND DONDALE: SPIDERS of NEWFOUNDLAND 


273 


TABLE 4. Proportions of Linyphiidae, Linyphiidae plus Lycosidae, and Linyphiidae plus Lycosidae plus Dictynidae in some 


northern spider faunas. 


BE?» QC SK? AB? MB? 
Total families 35 28 Zo 22 20 
Total species OF. OL], 550 a2 483 
Linyphiidae % 39,2 Ibo 36.0 2.) ee 
TL+L% LWA EG i) 43.5 46.9 45.3 
tL+L+D% 467. od 48.9 50,5. 49.7 


NE YR GL° BL’ HZ” a. Phe 
Figpe 15 10 8 4 3 
363 207 70 33 13 7 
32.6 45.8 64.3 69.7 69 re Wh 
2 WP 53.9 fom 81.8 84 85.7 
61.7 Bi Me) 78.6 84.9 aS, 100 


'Bennett 2001; Bennett et al. 2004*. * Bélanger and Hutchinson 1992, as revised by Paquin et al. 2001, with additions in 
Paquin and Dupérré 2003. *Buckle and Holmberg 2004*. *Aitchison-Benell and Dondale 1990. °Dondale et al. 1997. 
6Larsen and Scharff 2003. 7Koponen 1992. *Leech 1966. ’Braendegaard 1960. Some familial placements adjusted in accor- 


dance with Platnick 2005. 
+L+ L % = % of Linyphiidae + Lycosidae 
tL+L+D% = % of Linyphiidae + Lycosidae + Dictynidae 


three geographical attributes, total number of species, 
number of families and number of species within 
each family for each of the seven Canadian provinces 
or territories for which there is information. Locality 
columns are arranged left to right in order of most to 
least total species. There is a general correlation be- 
tween the three geographical attributes and the num- 
ber of families and species; analysis of the obvious 
exceptions will not be possible until knowledge of 
the faunas is standardised. 

Several authors (e.g. Leech 1966; Koponen 1992, 
1996; Larsen and Scharff 2003) have pointed out that 
northern spider faunas are numerically dominated by 
species of Linyphiidae, Lycosidae and Dictynidae in 
descending order of importance. For example, species 
of these three families make up 100% of the most 
northern spider fauna in the world on Peary Land, 
Greenland (c. 82.5°N), excluding for present purpos- 
es the introduced agelenid Tegenaria domestica 
(Braendegaard 1960). Table 4 compares the cumula- 
tive proportions of these three families (i.e. Linyphi- 
idae alone, Linyphiidae plus Lycosidae, Linyphiidae 
plus Lycosidae plus Dictynidae) as a scale of north- 
ernness for the seven Canadian areas considered in 
Table 3. In addition Table 4 includes the proportions 
in two other Canadian locations, the Belcher Islands 
(56° N) and Hazen Camp (82° N), and two non- 
Canadian locations, Peary Land (82.5° N) and the 
whole of Greenland (60°-82.5° N). The overall trend 
| of increasing numerical dominance of Linyphiidae, 
| Lycosidae and Dictynidae is confirmed. 
| Other Canadian spider fauna surveys have includ- 
| ed species likely to be discovered in the area in ques- 
| tion in the future (e. g. Dondale et al. 1997; Bélanger 
| and Hutchinson 1992). While this may be justified 
| when species absent from the area in question have 
| been reported from a contiguous land mass, it seems 
less useful when dealing with an island. 


Acknowledgments 

We thank Lloyd Hollett and Kevin Pardy who 
donated many Newfoundland spiders to the CNC. 
We are grateful for access to the valuable collections 
made by Phil Taylor and his team (Acadia Universi- 
ty), for the collections made by Anne-Marie Hynes 
and Shelley Pardy, and for the funding for the latter 
collections provided by the Newfoundland Inland 
Fish & Wildlife Division. We thank Parks Canada for 
permission to collect in Gros Morne National Park 
over several years. We are grateful to Dave Larson 
for providing access to his arthropod collections. We 
would also like to acknowledge all other collectors, 
too numerous to mention individually, who have con- 
tributed Newfoundland specimens directly or indirect- 
ly to the CNC. We are grateful to Robb Bennett, Don 
Buckle and Robert Holmberg for permission to use 
unpublished data. We thank Jim Redner for his many 
hours of labour on Newfoundland specimens. We 
want to thank the Biology Department, Memorial 
University, for facilities and supplies. JRP thanks Dave 
Larson for his encouragement and advice. 


Documents Cited (marked in text*) 

Bennett, R. G., D. L. Blades, D. Buckle, C. D. Dondale 
and R. C. West 2004. Unpublished data. Personal com- 
munication R. G. Bennett. 

Buckle, D. J., and R. G. Holmberg. 2004. Unpublished 
data. Personal communication D. J. Buckle. 

Natural Resources Canada: Canadian Geographical 
Names. 2005 http://gnss.nrcan.gc.ca/gnss-srt/searchName. 
jsp’?language=en 

Government Website, Alberta. 2004. www.gov.ab.ca 

Government Website, British Columbia. 2004. www.gov. 
be.ca 

Government Website, Manitoba. 2004. www.gov.mb.ca 

Government Website, Newfoundland and Labrador. 2004. 
www.gov.nf.ca 

Government Website, Quebec. 2004. www.gouv.qe.ca 


274 


Government Website, Saskatchewan. 2004. www.gov.sk.ca 

Government Website, Yukon. 2004. www.gov.yk.ca 

Platnick, N. I. 2005. The world spider catalog, version 5.5. 
American Museum of Natural History, online at http:// 
research.amnh.org/entomology/spiders/catalog/ 


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Received 21 June 2004 
Accepted | June 2005 


Additions and Range Extensions to the Vascular Plant Flora of the 
Continental Northwest Territories and Nunavut, Canada III 


WILLIAM J. Copy! and KENNETH L. READING? 


‘Biological Resources Program, Biodiversity, National Program on Environmental Health, Agriculture and Agri-Food Canada, 
Wm. Saunders Building (49), Central Experimental Farm, Ottawa, Ontario K1A 0C6, Canada 
11 Colborne Street, Thornhill, Ontario L3T 1Z4, Canada 


Cody, William J,. and Kenneth L. Reading. 2005. Additions and range extensions to the vascular plant flora of the continental 
Northwest Territories and Nunavut, Canada III. Canadian Field-Naturalist 119(2): 276-290. 


Fifteen native taxa are reported as new to either District of Keewatin (now Nunavut in part) or District of Mackenzie (Nunavut 
contains the northeastern portion): Aster puniceus, Astragalus eucosmus, Astragalus eucosmus f. albinus, Carex media, Epilobium 
latifolium f. albiflorum, Eriophorum viridi-carinatum, Erysiumum pallasii, Moneses uniflora, Najas flexilis, Potamogeton obtu- 
sifolius, Salix arctica x glauca, Salix glauca ssp. callicarpae, Salix myrtillifolia, Salix rotundifolia, Shepherdia canadensis, 


taxa are new to Northwest Territories overall. Significant range extensions for 157 native taxa are included. 


Key Words: Vascular plants, Mackenzie, Keewatin, Nunavut, flora, new records, range extensions, phytogeography. 


Since the publication of Additions and Range Exten- 
sions to the Vascular Plant Flora of the Continental 
Northwest Territories and Nunavut, Canada II (Cody 
et al. 2003) a large number of specimens collected by 
Kenneth L. Reading in the Continental Northwest Ter- 
ritories between the years of 1978 and 1995 were dis- 
covered and forwarded to the senior author for exami- 
nation. Many of these proved to be significant records 
and are reported here along with some additional speci- 
mens collected by D. Bush, D. Campbell, B. Cornish, V. 
J. Kajima, L. Kershaw, J. B. Korol, J. Lancaster, J. V. Mat- 
thews, J. W. Thieret, R. J. Reich, and J. Thompson. 

All of these specimens are preserved in the Agricul- 
ture and Agri-Food Canada Herbarium (DAO). As in 
Cody et al (2003) the authors have continued to use 
the historic names “District of Keewatin” and “District 
of Mackenzie” to follow the format of Porsild and Cody 
(1980) Vascular Plants of Continental Northwest Terri- 
tories and subsequent publications. However, continen- 
tal “Keewatin” and the former northeastern portion of 
“District of Mackenzie” are now part of the recently 
defined Continental Nunavut Territory. 

As in paper I a synopsis of the taxa addressed in 
the body of this paper follows with species listed in 
alphabetical order within categories. The taxa are then 
discussed in an annotated list by family in the same 
order as presented in the Flora of the Continental North- 
west Territories (Cody and Porsild 1980) together with 
citation of specimens and other pertinent information. 


New Taxa to the Continental Northwest 
Territories Status 

New Taxa to the Continental Northwest Territories 
(Keewatin): (10) 

Astragalus eucosmus 

Astragalus eucosmus f. albinus 

Carex media 

Epilobium latifolium f. albiflorum 


Eriophorum viridi-carinatum 
Erysimum pallasii 

Moneses uniflora 

Salix glauca ssp. callicarpaea 
Salix myrtillifolia 

Shepherdia canadensis 


New Taxa to the Continental Northwest Territories 
(Mackenzie): (5) 

Aster puniceus 

Potamogeton obtusifolius 

Najas flexilis 

Salix arctica X glauca 

Salix rotundifolia 


Range Extensions of Native Taxa in the Continental — 
Northwest Territories (Keewatin): (48) i 
Anemone parviflora 

Anemone richardsonii 
Antennaria isolepis 

Caltha natans 

Campanula uniflora 

Cardamine bellidifolia 
Cardamine pratensis 

Carex canescens 

Carex rariflora 

Carex rotundata 

Carex scirpoidea 

Cystopteris fragilis 

Draba fladnizensis 

Draba nivalis 

Draba wahlenbergii 

Dryopteris fragrans 

Epilobium davuricum 
Equisetum palustre 

Festuca brachyphylla 

Galium trifidum 

Juncus arcticus 

Juncus castaneus ssp. castaneus 
Ledum groenlandicum 
Minuartia biflora 

Minuartia rossii 


276 


2005 CopDY AND READING: VASCULAR FLORA OF NWT AND NUNAVUT 


Minuartia rubella 
Polygonum viviparum 
Potentilla nivea ssp. nivea 
Potentilla palustris 

Pyrola secunda 

Ranunculus sabinei 
Ranunculus x spitzbergensis 
Rubus arcticus ssp. acaulis 
Rubus idaeus 

Sagina caespitosa 

Salix arctophila 

Salix fuscescens 

Salix planifolia 

Salix reticulata 

Salix richardsonii 

Saxifraga nelsoniana ssp. porsildiana 
Saxifraga rivularis 

Scirpus caespitosus ssp. austriacus 
Silene walensis 

Stellaria calycantha 
Stellaria longipes 

Tofieldea coccinea 

Tofieldia pusilla 


Range Extensions of Native Taxa in the Continental 
Northwest Territories (Mackenzie): (109) 


Alisma triviale 

Anemone parviflora 
Arctostaphylos alpina 

Arnica angustifolia ssp. attenuata 
Arnica chamissonis 

Artemisia furcata 

Artemisia tilesii 

Aster pygmaeus 

Aster sibiricus 

Astragalus australis 

Braya glabella 

Braya humilis 

Calamagrostis lapponica 
Calamagrostis purpurascens 
Calamagrostis stricta ssp. stricta 
Callitriche hermaphroditica 
Caltha palustris var. arctica 
Cardamine bellidifolia 

Carex bebbii 

Carex nardina 

Carex scirpoidea 

Carex subspathacea 

Carex supina ssp. spaniocarpa 
Carex vaginata 

Cerastium alpinum 

Cerastium beeringianum 
Chenopodium dessicatum 
Chrysosplenium tetrandrum 
Crepis nana 


Cryptogramma crispa var. acrostichoides 


Cystopteris fragilis 
Cystopteris montana 
Danthonia spicata 
Descurainia sophioides 
Draba juvenalis 

Draba wahlenbergii 
Dryopteris fragrans 
Elymus canadensis 
Elymus macrourus 


Elymus trachycaulus ssp. violaceus 
Epilobium palustre 

Equisetum palustre 

Erigeron uniflorus ssp. eriocephalus 
Eriophorum angustifolium 
Eriophorum callitrix 
Eriophorum triste 

Eriophorum vaginatum 
Erysimum pallasii 

Eutrema edwardsii 

Festuca richardsonii 

Gentiana affinis 

Hedysarum alpinum 

Hedysarum boreale ssp. mackenzii 
Hierochloe hirta ssp. arctica 
Juncus arcticus 

Juncus castaneus ssp. castaneus 
Juncus stygius ssp. americanus 
Juncus triglumis ssp. albescens 
Juniperus communis 

Lagotis glauca 

Limosella aquatica 

Linnaea borealis var. americana 
Lupinus arcticus 

Luzula wahlenbergii 
Lycopodium selago 

Lycopus uniflorus 

Matricaria ambigua 

Minuartia yukonensis 
Muhlenbergia glomerata 
Myriophyllum alterniflorum 
Nymphaea tetragona ssp. leibergii 
Oxytropis arctobia 

Oxytropis deflexa ssp. foliosa 
Oxytropis maydelliana 

Papaver radicatum 

Parrya arctica 

Pedicularis capitata 

Pedicularis langsdorfii ssp. arctica 
Pedicularis lapponica 
Pedicularis macrodonta 
Phragmites australis 

Poa alpina 

Poa glauca 

Poa pratensis ssp. alpigena 
Potamogeton filiformis 
Potamogeton foliosus 

Potentilla biflora 

Ranunculus aquatilis var. eradicatus 
Rosa blanda 

Rhynchospora alba 

Salix alaxensis 

Salix arctica 

Salix glauca var. acutifolia 

Salix planifolia 

Salix reticulata 

Salix richardsonii 

Salix sphenophylla 

Saxifraga nivalis 

Scirpus rollandii 

Senecio congestus 

Silene walensis 

Solidago graminifolia var. major 
Sparganium hyperboreum 
Spartina gracilis 


277 


278 


Stellaria longipes 

Triglochin maritimum 

Trisetum spicatum 

Vaccinium vitis-idaea var. minus 
Woodsia ilvensis 


Annotated List 


LYCOPODIACEAE 
Lycopodium selago L., Mountain Club-moss — 
MACKENZIE: Hope Bay Area, northeast of Bathurst 
Inlet, Windy Lake, 68°03'15"N 106°37'00"W, K. 
Reading 152, 14 July 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 175 kilometers north of a site 
adjacent to the Arctic Circle (Porsild and Cody 1980). 


EQUISETACEAE 

Equisetum palustre L., Marsh Horsetail — MACKENZIE: 
muddy tundra pond margin, Coppermine River Area, 
67°31'15"N 116°01'00"W, K. Reading 8, 31 Aug. 2002 
(DAO); rocky fen pools, Coppermine River Area, 
67°33'30"N 116°14'54"W, K. Reading 47,21 Aug. 2002 
(DAO); muddy flooded rock polygon, Coppermine River 
Area, 67°34'00"N 116°17'00"W, K. Reading 9, | Sept. 
2002 (DAO); Hope Bay Area, northeast of Bathurst 
Inlet, Windy Lake, 68°03'15"N 106°37'00"W. K. Read- 
ing 204, 16 July 1987 (DAO); KEEWATIN: SY Area, 
62°12'20"N 97°52'00"W, K. Reading 121, 17 Aug. 1984 
(DAO). 

The first four specimens cited above extend the known 
range of this species about 500 kilometers northeast of the east 
end of Great Bear Lake (Porsild and Cody 1980). The fifth 
specimen is about 170 kilometers south of the only previously 
known localities in central KEEWATIN (Cody et al. 2003). 


PTERIDACEAE 
Cryptogramma crispa (L.) R.Br. var. achrostichoides 
(R.Br.) C.B. Clarke, Mountain-parsley — MACKENZIE: 
common on outcrop, 63°00'00”N 109°55'00"W, K. 
Reading 426, 8 Aug. 1995 (DAO). 

The specimen cited above is an extension of the known 


range in the Territory of about 85 kilometers northeast of the 
east end of Great Slave Lake (Porsild and Cody 1980). 


ASPIDIACEAE 
Cystopteris fragilis (L.) Bernh., Fragile Fern — 
MACKENZIE: 63°00'00"N_ 109°55'00”W, K. Reading 
423, 11 Aug. 1995 (DAO); KEEwaTIN: Griffin Lake, 
61°17'20"N 98°42'00”"W, K. Reading 98, 19 Aug. 
1990 (DAO). 

The first specimen cited above is an extension of the 
known range in the Territory of about 70 kilometers north- 
east of the east end of Great Slave Lake and the second 
specimen is the southernmost yet known in the District of 
Keewatin (Porsild and Cody 1980). 


Cystopteris montana (Lam.) Bernh., Mountain Bladder 
Fern — MACKENZIE: uncommon under tamaracks, 
63°00'00"N 109°52'00"W, K. Reading 429, 9 Aug. 
1995 (DAO). 

This is a rare species in the Continental Northwest Terri- 
tories which Porsild and Cody (1980) knew from only three 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


localities west of the west end of Great Slave Lake. The 
specimen cited above is an extension of the range to the east 
of Great Slave Lake of about 390 kilometers. 


Dryopteris fragrans (L.) Schott, Fragrant Cliff Fern — 
MACKENZIE: Hope Bay Area, northeast of Bathurst 
Inlet, Roberts Lake, 68°10'30"N 106°33'00"W, K. Read- 
ing 231, 11 Aug. 1987 (DAO); KEEwaTIN: south south- 
west of Bissett Lake, 63°44'00"N 95°30'00"W, K. Read- 
ing 336, 28 Aug. 1982 (DAO). 

This is a widespread species in Canada. The first specimen 
cited above is an extension of the known range of about 180 
kilometers north northwest from the Arctic Circle and the 


second specimen is from a site between two sites south of 
the Arctic Circle (Porsild and Cody 1980). 


Woodsia ilvensis (L.) R.Br., Rusty Woodsia — MAc- 
KENZIE: 63°00'00"N 109°55'00" W, K. Reading 425, 8 
Aug. 1995 (DAO); outcrop beside bog, 62°59'00"N 
109°51'00"W, K. Reading 428, 9 Aug. 1995 (DAO); 
Hope Bay area, northeast of Bathurst Inlet, Roberts 
Lake, 68°10'30"N 116°33'00"W, K. Reading 213, 11 
Aug. 1987 (DAO). 

The first two specimens cited above are an extension of the 
known range in the Territory (Porsild and Cody 1980) of 
about 170 kilometers northeast of a site near the east end of 
Great Slave Lake. The second specimen is a northward 


extension of about 75 kilometers from a site reported by Cody 
et al. (1984). 


CUPRESSACEAE 

Juniperus communis L. s.1., Ground Juniper — MACKENZIE: 
dolomite scarp, Coppermine River Area, 67°09'00"N 
115°45'00"W, K. Reading 83, 28 Aug. 2002 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers northeast of a 
site at the east end of Great Bear Lake (Porsild and Cody 
1980). 


SPARGANIACEAE 
Sparganium hyperboreum Laest. — MACKENZIE: big 
pool in creek, Coppermine River Area, 67°31'10"N 
116°10'05"W, K. Reading 52, 22 Aug. 2002 (DAO). 
The specimen cited above is from a site midway between 
a site north of the east end of Great Bear Lake and the south 
end of Bathurst Inlet (Porsild and Cody 1980). This species 
was only found once in this area. 


POTAMOGETONACEAE 
Potamogeton filiformis Pers., Fine-leaved Pondweed 
— MACKENZIE: sandy Lake, Coppermine River Area, 
67°06'05"N_ 115°44'50"W, K. Reading 50, 21 Aug. 
2002 (DAO). 

The specimen cited above is from a site midway between 
a site north of the east end of Great Bear Lake and Bathurst 
Inlet (Porsild and Cody 1980). 


Potamogeton foliosus Raf. (P. foliosus Raf. var. macel- 
lus Fern.), Closed-leaved Pondweed — MACKENZIE: 
61.7382°N 121.1534°W, L. Kershaw 18, 2003 (DAO). 

This species was considered rare in the Northwest Terri- 
tories by McJannet et al. (1995), who knew it only from the 
vicinities of Yellowknife, Nahanni National Park and the 
Great Bear River areas. The specimen cited above is from a 
site in the Keller Lake area east of the Mackenzie River. 


2005 


Potamogeton obtusifolius M.&K., Blunt-leaved 
Pondweed — MACKENZIE: Mackenzie River- Yellowknife 
Highway, small lake at Mile 12.7 S, Thieret & Reich 
8358, 28 July 1961 (DAO); same area, small lake, Mile 
38.3 S, Thieret & Reich 4947, 23 July 1961 (DAO); 
62.158°N 122.4891°W, L. Kershaw 19, 2003 (DAO). 

Porsild and Cody (1980) suggested that this species was 
to be expected in the southern parts of the Precambrian 
Shield area. Haynes (1974) reported it from the vicinity of 
Yellowknife. Haynes had already revised the first three 
specimens in 1971, but this revision was overlooked by Por- 
sild and Cody. In 1995 he confirmed his identification. The 
fourth specimen cited above from a site near the Mackenzie 
River is an extension of the known range in the Territory of 
about 400 kilometers to the west. 


NAJADACEAE 
Najas flexilis (Willd.) Rostk. & Schmidt, Wavy Water 
Nymph — MACKENZIE: beside beaver lodge near north- 
west corner of unnamed lake, 62.9854°N 123.1753°W, 
L. Kershaw 14, 2003 (DAO). 

The specimen cited above is new to the flora of the Con- 
tinental Northwest Territories (Porsild and Cody 1980). 


SCHEUCHZERIACEAE 
Triglochin maritimum L., Seaside Arrow-grass — MAC- 
KENZIE: wet fen, Coppermine River Area, 67°34'10"N 
116°18'SO"W, K. Reading 153, 1 Sept. 2002 (DAO). 
The specimen cited above is from a location midway 
between a site at the northeast end of Great Bear Lake and a 
site near the south end of Bathurst Inlet (Porsild and Cody 
1980). 


ALISMATACEAE 
Alisma triviale Pursh (A. plantago-aquatica L. vat. 
americanum J. A. Schultes) — MACKENZIE: Willowlake 
River, 62.7024°N 123.1043°W, sand and gravel bar 
on the north side of the river, D. Soppet 2, 2003 (DAO). 
This species was considered rare in the Territory on the 
basis of a single collection from the shoreline of Hanging 
Ice River, north of Fort Smith (McJannet et al. 1995). The 
specimen cited above is an extension of the known range in 
the Territory of about 500 kilometers to the northwest. 


POACEAE 


Calamagrostis lapponica (Wahlenb.) Hartm. — MAc- 


KENZIE: Camp Peninsula on September (Mouse) Lake, 
Coppermine River Area, 67°06'05"N_ 115°44'SO"W, 
K. Reading 115, 24 Aug. 2002 (DAO). 

The specimen cited above is from a site midway between 
the east end of Great Bear Lake and the south end of Bathurst 
Inlet (Porsild and Cody 1980). 


Calamagrostis purpurascens R.Br. — MACKENZIE: Hope 
Bay Area, northeast of Bathurst Inlet, Windy Lake, 
68°03'1S"N 106°37'00"W, K. Reading 318, 29 June 
1987 (DAO). 

The specimen cited above is from a site about 170 kilo- 
meters north of the south end of Bathurst Inlet and is the 


most northwestern yet known from the District of Mackenzie 
(Porsild and Cody 1980). 


Calamagrostis stricta (Timm) Koeler ssp. stricta (C. 
neglecta (Ehrh.) Gaertn., Mey. & Scherb.) — MACKENZIE: 


Copy AND READING: VASCULAR FLORA OF NWT AND NUNAVUT 


299 


“Lunch outcrop”, Coppermine River Area, 67°32'15"N 
116°02'30"W, K. Reading 110, 26 Aug. 2002 (DAO); 
Hope Bay Area, northeast of Bathurst Inlet, Windy 
Lake, 63°03'15"N 106°37'00"W, K. Reading 304, 15 
July 1978 (DAO). 

The first specimen cited above is from a site about 250 
kilometers northeast of the east end of Great Bear Lake 
(Porsild and Cody 1980). To the north on Victoria Island it 
is known about 250 kilometers away, and to the east about 
630 kilometers. The second specimen is from a site about 
290 kilometers northeast of the first specimen location. 


Danthonia spicata (L.) Beauv., Poverty Oatgrass — 
MACKENZIE: along break between the sandbar and the 
Black Spruce forest on the edge of the river, north side 
sandbar of A River between Two Mountains, 62.942°N 
123.0399°W, D. Soppet 6, 2003 (DAO). 

Porsild and Cody (1980) suggested that this species 
should be looked for along the southern rim of the Precam- 
brian Shield east of the Slave River. Specimens which were 
all collected in Nahanni National Park and identified as D. 
intermedia have since been revised to D. spicata. The speci- 
men cited above is from a site northeast of the park. 


Elymus canadensis L., Canadian Wild Rye — MACKEN- 
ZIE: upper shore of Mackenzie River about 100 m north 
of ferry landing, 62.1498°N 122.5327°W, L. Kershaw 
7, 2003 (DAO). 

This is a rare species in the Territory (McJannet et al. 1995). 
The specimen cited above is an extension of the known 
range of about 175 kilometers north of a site adjacent to the 
Liard River. It is however not far distant from a site on the 
Norman Wells Pipeline adjacent to the Willowlake River 
winter road which presumably was introduced (Cody et al. 2000). 


Elymus macrourus (Turez.) Tzvelev (Agropyron seri- 
ceum Hitchc.) — MACKENZIE: Hope Bay Area, northeast 
of Bathurst Inlet, Windy Lake, 68°03'15"N 106°37'00"'W, 
K. Reading 310, 29 June 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 750 kilometers east of sites 
adjacent to the Arctic Coast (Porsild and Cody 1980). 


Elymus trachycaulus (Link) Gould ex Shinners ssp. vio- 
laceus (Hornem.) A. & D. Léve (Agropyon trachy- 
caulum (Link) Malte sl.) — MACKENZIE: sand ridge, 
Coppermine River Area, 67°33'00"N 115°58'00"W, 
K. Reading 97A, 5 Sept. 2002 (DAO) (determined by 
S. Darbyshire 2002). 

The specimen cited above is an extension of the known 
range of this group (Porsild and Cody 1980) of about 250 kilo- 
meters northeast of the east end of Great Bear Lake. 


Festuca brachyphylla Schultes — KEEWATIN: Bray Lake, 
61°26'N 97°59'W, K. Reading 104, 21 June 1990 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 150 kilometers to the northeast 
of a site in the extreme southwest of the Territory (Porsild 
and Cody 1980). 


Festuca richardsonii Hook. (F- rubra L. ssp. richard- 
sonit (Hook.) Hultén) — MACKENZIE: Hope Bay Area, 
northeast of Bathurst Inlet, Windy Lake, 68°03'15"N 
106°37'00"W, K. Reading 312B, 29 June 1987 (DAO). 


280 


The specimen cited above is an extension of the known 
range in the Territory of about 200 kilometers north of the 
south end of Bathurst Inlet (Porsild and Cody 1980). 


Hierochloe hirta (Schrank) Borbas ssp. arctica (G.) 
Weim. (H. odorata pro parte sensu Porsild and Cody 
(1980), Sweet Grass — MACKENZIE: Hope Bay Area, 
northeast of Bathurst Inlet, Windy Bay, 68°03'15S"N 
106°37'00"W, K. Reading 317, 29 June 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 540 kilometers northeast of the 
northeast end of Great Bear Lake (Porsild and Cody 1980). 


Muhlenbergia glomerata (Willd.} Trin. — MACKENZIE: 
mineral spring, Mackenzie Valley pipeline survey, 
61°37'17"N 121°30'14'"W, L. Kershaw s.n., 8 Aug. 2002 
(DAO). 

The specimen cited above which is the northernmost yet 
found in the Territory is an extension of the known range of 
about 300 kilometers to the northwest of a site adjacent to 
the Liard River mapped by Porsild and Cody (1980). 


Phragmites australis (Cav.) Trin. ex Steud., (P. com- 
munis Trin. var. berlandieri (Fourn.) Fern.), Common 
Reed — MACKENZIE: shallow water about 10 m from the 
shore of a large alkaline lake, 61.7617°N 121.0793°W, 
L. Kershaw 17, 2003 (DAO). 

The specimen cited above from the vicinity of Fort Simp- 
son is from only the second known site in the Territory about 
130 kilometers to the northeast in the vicinity of Yohin Lake 
(Scotter and Cody 1974). 


Poa alpina L., Alpine Blue Grass — MACKENZIE: Cop- 
permine River Area, Camp Peninsula on September 
(Mouse) Lake, 67°06'05"N 115°44'50"W, K. Reading 
118, 24 Aug. 2002 (DAO); Hope Bay Area, northeast 
of Bathurst Inlet, Windy Lake, 68°03'15"N 106°37'00"'W, 
K. Reading 321, 29 June 1987 (DAO). 

The first specimen cited above extends the known range 
in the Territory about 125 kilometers west and the second 
specimen about 125 kilometers north of a site adjacent to 
Burnside Landing at latitude 66°51'N adjacent to Bathurst 
Inlet (Cody et al. 1984). 


Poa glauca Vahl — MACKENZIE: Hope Bay Area, northeast 
of Bathurst Inlet, Windy Lake, 68°03'15"N 106°37'00"W, 
K. Reading 313, 29 June 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers from the south 
end of Bathurst Inlet (Porsild and Cody 1980). 


Poa pratensis L. ssp. alpigena (L.) Lindm. (P. alpige- 
na (Fr.) Lindm.) — MACKENZIE: George Lake Camp, 
65°53'10"N 107°23'00"W, K. Reading 463, 467, 468, 
469, 16 Aug. 1988 (DAO); Hope Bay Area, northeast 
of Bathurst Inlet, Windy Lake, 68°03'05"N 106°37'00"W, 
K. Reading 316, 319, 29 June 1987 (DAO). 

The first location cited above is about 85 kilometers south 
of sites near latitude 67°N reported by Cody et al. (1984). The 
Windy Lake sites are the northeasternmost yet known from 


the Territory and are about 130 kilometers north of latitude 
67°N. 


Spartina gracilis Trin., Alkali Cord Grass — MACKENZIE: 
sandbar on north side of river, Willowlake River, 
62.7024°N 123.1043°W, D. Soppet 28, 2003 (DAO). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


This is a rare species in the Territory (McJannet et al. 1995). 
The specimen cited above is an extension of the known range 
of about 240 kilometers northwest of the vicinity of Fort 
Simpson. 


Trisetum spicatum (L.) Richt. - MACKENZIE: George 
Lake Camp, 65°53'10"N 107°23'8"W, K. Reading 470, 
18 Aug. 1988 (DAO); Hope Bay Area, northeast of 
Bathurst Inlet, Windy Lake, 68°03'15"N 106°37'00"W, 
K. Reading 306, 312A, 320, 322, 29 June 1987 (DAO). 
The first location cited above is about 85 kilometers south 
of sites near latitude 67°N reported by Cody et al. (1984). 
The Windy Lake site is the northeasternmost yet known from the 
Territory and is about 130 kilometers north of latitude 67°N. 


CYPERACEAE 
Carex bebbii Olney — MACKENZIE: Mackenzie River 
Valley pipeline survey, steep boulder strewn stream 
shore of Mackenzie River near Liard, 61°50'53"N 
121°10'7"W, L. Kershaw s.n., 8 Aug. 2002 (DAO). 
Although widespread across much of Canada, Porsild and 
Cody (1980) knew this rare plant in the Territory from only 
four locations: near Fort Smith, Fort Good Hope, Fort Simp- 
son and Fort Liard. 


Carex canescens L. — KEEWATIN: Griffin Lake, 
61°16'30"N 98°42'00"W, K. Reading 100, 26 July 1990 
(DAO). 

The specimen cited above is the northernmost yet found 
in the Territory and is from a site about 60 kilometers north- 
east of a site mapped by Porsild and Cody (1980). 


Carex media R.Br. — KEEWATIN: Griffin Lake, 
61°16'30"N 98°42'00"W, K. Reading 99, 26 July 1990 
(DAO). 

The specimen cited above is the first known from this 
Territory. It is from a site about 85 kilometers northeast of a 


site in the province of Manitoba, just south of latitude 60°N 
(Porsild and Cody 1980). 


Carex nardina Fries — MACKENZIE: Hope Bay Area, 
northeast of Bathurst Inlet, Windy Lake, 68°03'15"N 
106°37'00"W, K. Reading 301, 14 July 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers northeast of a 
location cited by Cody et al. (1984) and is the northeasternmost 
yet known in the Territory. 


Carex rariflora (Wahlenb.) Sm. — KEEWATIN: west of 
Imikula Lake, 62°10'00"N 97°40'20"W, K. Reading 
397, 10 Aug. 1983 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 80 kilometers inland from sites 
adjacent to the Hudson Bay coast (Porsild and Cody 1980). 


Carex rotundata Wahl. — KEEWATIN: SY _ area, 
62°12'20"N 97°53'00"W, K. Reading 219, 19 July 1984 
(DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 80 kilometers northeast of a 
site in the extreme south mapped by Porsild and Cody (1980). 


Carex scirpoidea Michx. — MACKENZIE: sand flats, 
Coppermine River area, 67°06'05"N_ 115°44'50"W, 
K. Reading 15A, 8 Sept. 2002 (DAO); Hope Bay Area, 
northeast of Bathurst Inlet, Windy Lake, 68°03'15"N 


2005 


106°37'00"W, K. Reading 287, 15 July 1987 (DAO); 
KEEWATIN: Griffin Lake, 61°16'30"N 98°42'00"W, K. 
Reading 226, 26 July 1990 (DAO). 

The first specimen cited above is from a site midway 
between Coppermine and the south end of Bathurst Inlet 
(Porsild and Cody 1980). The second specimen is an exten- 
sion of the known range in the Territory of about 170 kilo- 
meters northeast of the south end of Bathurst Inlet and is the 
northeasternmost yet known in the Territory. The third spec- 
imen is an extension of about 100 kilometers northeast of 
the southwesternmost previously known site in Keewatin 
(Porsild and Cody 1980). 


Carex subspathacea Wormsk}j. — MACKENZIE: Hope 
Bay Area, northeast of Bathurst Inlet, Windy Lake, 
68°03'1S"N 106°37'00"W, K. Reading 283, 15 July 
1987 (DAO). 

The specimen cited above of this rare northern species is 
from a site about 170 kilometers northeast of the south end 
of Bathurst Inlet (Porsild and Cody 1980) and is the east- 
ernmost yet found in the Territory. 


Carex supina Wahl. ssp. spaniocarpa (Steud.) Hultén 
— MACKENZIE: Hope Bay Area, northeast of Bathurst 
Inlet, Windy Lake, 68°03'15"N 106°37'00"W, K. Read- 
ing 311, 29 June 1987 (DAO). 

The specimen cited above is from a site about 170 kilo- 
meters northeast of the south end of Bathurst Inlet (Porsild 
and Cody 1980) and is the northeasternmost yet found in the 
Territory. 


Carex vaginata Tausch — MACKENZIE: Hope Bay Area, 
northeast of Bathurst Inlet, Roberts Lake, 68°10'30"N 
106°33'00"W, K. Reading 330, 4 Aug. 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 150 kilometers northeast of a 
site near Burnside Landing adjacent to Bathurst Inlet (Cody 
et al. 1984). 


Eriophorum angustifolium Honckn. — MACKENZIE: 
Hope Bay Area, northeast of Bathurst Inlet, Windy Lake, 
68°03'15"N 106°37'00"W, K. Reading 202, 15 July 1987 
(DAO); Hope Bay Area, northeast of Bathurst Inlet, 
Roberts Lake, 68°10'30"N 106°33'00"W, K. Reading 
261, 1 Aug. 1987 (DAO). 

The specimens cited above are a northern extension of 
the known range in the Territory of about 170 kilometers 


north of a site east of the south end of Bathurst Inlet (Por- 
sild and Cody 1984). 


Eriophorum callitrix Cham. — MACKENZIE: George 
Lake Camp, 65°53'12"N 107°23'8"W, K. Reading 465, 
18 Aug. 1988 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 80 kilometers south of the south 
end of Bathurst Inlet (Porsild and Cody 1980). 


Eriophorum triste (Th. Fr.) Hadac & Love — MACKENZIE: 
Hope Bay Area, northeast of Bathurst Inlet, Windy Lake, 
68°03'15"N 106°37'00"W, K. Reading 286, 5 July 1987 
(DAO). 

The specimen cited above of this circumpolar arctic- 
alpine species is an extension of the known range in the Ter- 
ritory of about 500 kilometers northeast of a site adjacent to 
the east end of Great Bear Lake. Adjacent to the Arctic 


CopY AND READING: VASCULAR FLORA OF NWT AND NUNAVUT 


281 


Coast it is known about 400 kilometers to the west and it is 
also known from Victoria Island (Porsild and Cody 1980). 


Eriophorum vaginatum L. — MACKENZIE: Hope Bay 
Area, northeast of Bathurst Inlet, Windy Lake, 
68°03'15"N 106°37'00"W, K. Reading 285, 5 July 
1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 150 kilometers northeast of sites 
adjacent to the middle of Bathurst Inlet (Cody et al. 1984). 


Eriophorum viridi-carinatum (Engelm.) Fern. — KEE- 
WATIN: SY area, 62°12'30"N 97°55'00"W, K. Reading 
252, 17 Aug. 1984 (DAO). 

This species is new to Continental District of Keewatin. It 
is known to the west in the District of Mackenzie about 625 
kilometers away at the east end of Great Slave Lake and to 
the southwest in Saskatchewan about 475 kilometers away 
just south of the provincial border (Porsild and Cody 1980). 


Rhynchospora alba (L.) Vahl, White Beak-rush — MAc- 
KENZIE: 61.9954°N 121.5027°W, L. Kershaw 20, 2003 
(DAO). 

This is a rare species in the Northwest Territories (McJan- 
net et al. 1995) known from a single site near Hearst Lake 
(Cody and Talbot 1978) south of the east end of Great Slave 
Lake. The specimen cited above is from a site about 150 kilo- 
meters to the northwest. 


Scirpus caespitosus L. ssp. austriacus (Pallas) Asch. 
& Graebn. — KEEWATIN: east of Pebble Beach Lake, 
south of Yathkyed Lake, 62°22'00"N 97°28'00"W, K. 
Reading 178, 10 July 1985 (DAO). 

The specimen cited above is from a site midway between 
sites near the southern territories border and latitude 64°N 
(Porsild and Cody 1980). 


Scirpus rollandii Fern. — MACKENZIE: Mackenzie 
River Valley pipeline survey, mineral rich site near 
springs, mud hummocks, 65°32'24"N_ 127°36'47'"W, 
L. Kershaw s.n., 1 Aug. 2002 (DAO); hundred plus 
plants in calcareous area where stream widens and 
pools on flat area before flowing into mineral lake, 
Wrigley area, 62.7307°N 123.1098°W, D. Soppet 26A, 
2003 (DAO). 

McJannet et al. (1995) knew this rare species in the Terri- 
tory from only four localities, one near the northwest end of 
Great Slave Lake and three in the Mackenzie Mountains. The 
first specimen cited above is from a site northeast of the previ- 
ously northernmost site in the mountains. The second speci- 
men is from a site midway between the Mackenzie Moun- 
tain site and a site west of the north arm of Great Slave Lake. 


JUNCACEAE 

Juncus arcticus Willd. — MACKENZIE: Coppermine River 
Area, sandy lakeshore, 67°06’08"N 115°44'50"W, K. 
Reading 15B, 15 Aug. 2002 (DAO); KEEWATIN: Big Bird 
Lake, 62°17'30"N 97°38'00"W, K. Reading 180, 25 
June 1985 (DAO). 

The first specimen cited above is from a site about 100 kilo- 
meters west of Bathurst Inlet. The second specimen is from 
a site midway between a site adjacent to the Manitoba bor- 
der west of Hudson Bay and a site about latitude 63°30'N 
east of longitude 100°W (Porsild and Cody 1980). 


282 


Juncus castaneus Smith ssp. castaneus — MACKENZIE: 
Hope Bay Area, northeast of Bathurst Inlet, Roberts 
Lake, 68°10'30"N 106°33'00"W, K. Reading 329, 4 
Aug. 1987 (DAO); KEEWATIN: No-camp Lake, south 
of Yathkeyed Lake, 62°22'00"N 97°27'30"W, K. 
Reading 201, 21 July 1985 (DAO); Big Bird Lake, 
62°17'30"N 97°38'00"W, K. Reading 181, 25 June 
1985 (DAO). 

The first specimen cited above is an extension of the 
known range in the Territory of about 200 kilometers north- 
east of the south end of Bathurst Inlet. The second and third 
specimens are from sites about 125 kilometers north of a site 


in the interior of the District of Keewatin mapped by Porsild 
and Cody (1980). 


Juncus stygius L. ssp. americanus (Buch.) Hultén, Bog 
Rush — MACKENZIE: Mackenzie River Valley pipeline 
survey, sandy shoreline of creek, 61°19'27"N 
120°50'21"W, L. Kershaw s.n., 3 Aug. 2002 (DAO); 
in graminoid/shrub fen in lower wet spots, 60.6383°N 
120.3463°W, D. Soppet 9, 17 Aug. 2003 (DAO). 

The first specimen cited above from south of Fort Simpson 
is only the fourth known in the Territory. McJannet et al. 
(1995) knew it from just north of latitude 60°N from south 
of the east and west ends of Great Slave Lake and adjacent 
to Hudson Bay.This species was first recorded in the Terri- 
tory by J. W. Thieret (1959) on the basis of a specimen collect- 
ed by Thieret and Reich (1964) from among sedges at foot 
of “hummock island” in marly lake at mile 442 Enterprise- 
Mackenzie River Highway, about 215 kilometers east of the 
specimens cited above. 


Juncus triglumis L. ssp. albescens (Lange) Hultén (J. 
albescens (Lange) Fern.) —- MACKENZIE: Coppermine 
River Area, 67°06'05"N_ 115°44'50"W, K. Reading 
297, 3 Sept. 2002 (DAO). 

The specimen cited above is from a site midway between 
the south end of Bathurst Inlet and sites at the east end of 
Great Bear Lake (Porsild and Cody 1980). 


Luzula wahlenbergii Rupr. — MACKENZIE: George 
Lake Camp, 65°55'10"N_ 107°23'00"W, K. Reading 
466, 18 Aug. 1988 (DAO). 

The specimen cited above is from a site about 175 kilo- 
meters west of a site at about 101°W longitude just south of 
latitude 66°N (Porsild and Cody 1980). 


LILIACEAE 

Tofieldia coccinea Richards. — KEEWATIN: Kazan 
Falls, 63°43'00"N 95°51'30"W, K. Reading 348, 30 
Aug. 1982 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 150 kilometers east of speci- 
mens adjacent to longitude 100°W (Porsild and Cody 
1980). 


Tofieldia pusilla (Michx.) Pers. — KEEWATIN: south of 
Bissett Lake, 63°43'00"N 95°18'00"W, K. Reading 
347, 22 July 1982 (DAO). 

The specimen cited above is from a site about 150 kilo- 
meters east of a site just east of longitude 100°W (Porsild 
and Cody 1980). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


SALICACEAE 
Salix alaxensis (Anderss.) Cov. — MACKENZIE: sand flat 
below esker, Coppermine River Area, 67°33'00"N 
115°58'00"W, K. Reading 308, 5 Sept. 2002 (DAO) 
(determined by G. Argus). 

The specimen cited above is from a site midway between 


the south end of Bathurst Inlet and the vicinity of Coppermine 
(Porsild and Cody 1980). 


Salix arctica Pallas, Arctic Willow — MACKENZIE: Hope 
Bay Area, northeast of Bathurst Inlet, Windy Lake, 
68°03'15"N 106°37'00"W, K. Reading 442, 7 July 1987 
(DAO); Roberts Lake, 68°10'30"N 106°33'00"W, K. 
Reading 278, 24 June 1987, 250, 4 Aug. 1987, K. Read- 
ing 274, 10 Aug. 1987 (DAO) (determined by G. Argus). 

The four specimens cited above are a northeast extension 
of the known range in the Territory of about 170 kilometers 


from a site at the southern end of Bathurst Inlet (Porsild and 
Cody 1980). 


Salix arctica x glauca — MACKENZIE: Hope Bay Area, 
northeast of Bathurst Inlet, Windy Lake, 68°03'15"N 
106°37'00"W, K. Reading 448, 449, 450, 7 July 1987, 
14 July 1987 (DAO) (determined by G. Argus). 

This putative hybrid was not known to Porsild and Cody 
(1980). 


Salix arctophila Cockerell — KEEWATIN: southwest of 
Big Bird Lake, south of Yathkyed Lake, 62°16'50"N 
97°41'00"W, K. Reading 144, 28 June 1985 (DAO); 
southwest of Big Bird Lake, 62°17'00"N 97°40'00"W, 
K. Reading 30, 28 June 1985 (DAQO); southwest of 
No-camp Lake, south of Yathkyed Lake, 62°22'40"N 
97°27'30"W, K. Reading 436, 11 July 1985 (DAO); 
Camp 2, 61°21'00"N 97°53'30"W, K. Reading 447, 7 
July 1990 (DAO) (determined by G. Argus). 

The specimens cited above are from sites midway 


between the Hudson Bay Coast and a site just east of longi- 
tude 100°W (Porsild and Cody 1980). 


Salix fuscescens Anderss. — KEEWATIN: Big Bird Lake, 
62°18'00"N 97°39'00"W, K. Reading 36, 25 June 1985 
(DAO); Washaneepisuki Lake, 62°12'30"N 97°52'00"W, 
K. Reading 141, 17 July 1984 (DAO); north of No-camp 
Lake, 62°22'15"N 97°27'10"W, K. Reading 435, 11 July 
1985 (DAO) (determined by G. Argus). 

The specimens cited above are from sites midway between 
the Hudson Bay Coast and a site just east of longitude 100°W 
(Porsild and Cody 1980). 


Salix glauca L. var. acutifolia (Hook.) Schneid., 
Blue-green Willow — MACKENZIE: Hope Bay Area, 
northeast of Bathurst Inlet, Roberts Lake, 68°10'30"N 
106°33'00"W, K. Reading 276, 10 Aug. 1987 (DAO) 
(determined by G. Argus). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers northeast of the 
south end of Bathurst Inlet and is the northeasternmost site 
in the Territory (Porsild and Cody 1980). 


Salix glauca L. ssp. callicarpaea (Trautv.) Bocher — 
KEEWATIN: Bray Lake, 61°26'N 97°59'W, K. Reading 
451, 21 June 1990 (DAO); Washaneepisuki Lake, 
62°12'30"N 97°52'00"W, K. Reading 148, 17 July 1984 


2005 


(DAO); north of No-camp Lake, south of Yathkyed 
Lake, 62°22'15"N 97°28'10"W, K. Reading 138, 16 
July 1985 (DAO); north of No-camp Lake south of 
Yathkyed Lake, 62°22'00"N 97°30'05"W, K. Reading 
139, 22 July 1985 (DAO) (determined by G. Argus). 

This subspecies was known in northeastern North America 
from Greenland, Labrador, Newfoundland and Quebec (spec- 
imens in DAO determined by G. Argus). The specimens cited 
above extend the known range into central southern District 
of Keewatin. 


Salix myrtillifolia Anderss., Blueberry Willow — KEE- 
WATIN: Bray Lake, 61°26'N 97°59'W, K. Reading 444, 
21 June 1990 (DAO) (determined by G. Argus). 

This species is new to mainland District of Keewatin. The 
nearest site mapped by Porsild and Cody (1980) was in north- 
western Manitoba about 340 kilometers to the southwest and 
to the west in the District of Mackenzie it is common west of 
about longitude 106°W. 


Salix planifolia Pursh, Tea-leaf Willow — MACKENZIE: 
Hope Bay Area, northeast of Bathurst Inlet, Windy 
Lake, 68°03'15"N 106°37'00"W, K. Reading 446, 452, 
1 and 7 July 1987 (DAO); KEEWATIN: Washaneepisuki 
Lake, 62°12'30"N 97°52'00"W, K. Reading 142, 15 
July 1984 (DAO); Big Bird Lake, 62°17'30"N 
97°38'00"W, K. Reading 437, 25 June 1985 (DAO); 
62°20'00"N 97°22'00"W, K. Reading 32, 16 July 1985 
(DAO); Pebble Beach Lake south of Yathkyed Lake, 
62°22'00"N 97°30'00"W, K. Reading 149, 16 July 
1985 (DAO) (determined by G. Argus). 

The Windy Lake specimen cited above is an extension of 
the known range in the District of Mackenzie of about 170 
kilometers from sites adjacent to Bathurst Inlet south of lat- 
itude 67°N reported by Cody et al. (1984). The District of 
Keewatin sites are midway between the Hudson Bay Coast and 
about longitude 98°W. 


Salix reticulata L., Net-veined Willow — MACKENZIE: 
Hope Bay Area, northeast of Bathurst Inlet, Roberts 
Lake, 68°10'30"°N 106°33'00"W, K. Reading 279, 22 
July 1987 (DAO); KEEwaATIN: Camp 2, 61°21'00"N 
97°53'31"W, K. Reading 439, 7 July 1990 (DAO); 
Washaneepisuki Lake, west of Imikula_ Lake, 
62°12'30"N 97°52'00"W, K. Reading 145, 3 July 1984 
(DAO); Pebble Beach Lake south of Yathkyed Lake, 
62°22'00"N 97°30'00"W, K. Reading 146, 22 July 
1985 (DAO) (determined by G. Argus). 

The first specimen cited above is an extension of the 
known range in the District of Mackenzie of about 170 kilo- 
meters northeast of a site at the south end of Bathurst Inlet 
(Porsild and Cody 1980). The District of Keewatin speci- 
mens are from sites midway between the Hudson Bay Coast 
and longitude 101°W (Porsild and Cody 1980). 


Salix richardsonii Hook. (S. lanata L. ssp. richard- 
sonii (Hook.) Hultén, Richardson’s Willow — MACKEN- 
ZiIE: Hope Bay Area, northeast of Bathurst Inlet, 
Windy Lake, 68°03'15"N_ 106°37'00"W, K. Reading 
441, 445, 7 July 1987 (DAO); Hope Bay Area, north- 
east of Bathurst Inlet, Roberts Lake, 68°10'30"N 
106°33'W, K. Reading 275, 10 Aug. 1987 (DAO); 
KEEWATIN: Big Bird Lake, south of Yathkyed Lake, 


CopDY AND READING: VASCULAR FLORA OF NWT AND NUNAVUT 


283 


62°17'30"N 97°38'00"W, K. Reading 150, 25 June 
1985 (DAO); Pebble Beach Lake south of Yathkyed 
Lake, 62°22'00"N 97°52'00"W, K. Reading 151, 11 
June 1985 (DAO) (determined by G. Argus). 

The Mackenzie specimens cited above are an extension 
of the known range in the Territory of about 170 kilometers 
northeast of a site in the vicinity of Burnside Landing adja- 
cent to Bathurst Inlet. The Keewatin specimens extend the 
known range in the Territory about 250 kilometers north 
from a site adjacent to the Manitoba border (Porsild and Cody 
1980). 


Salix rotundifolia Trautv. ssp. rotundifolia, Round-leaf 
Willow — MACKENZIE: Mackenzie Mountains, lime- 
stone, stoney tundra slope, 17 miles NW Little Divide 
Lake, 63°17'N 128°17'W, W. J. Cody 16633, 26 July 
1967 (DAO); Mackenzie Mountains, in deep moss on 
granitic mountain, 5 miles SE of O’Grady Lake, 
62°S57'N 128°58'W, W. J. Cody 16750, 27 July 1967 
(DAO); Mackenzie Mountains, 10 miles NE of 
O’Grady Lake, 63°0S'N 128°50'W, W. J. Cody 16889, 
29 July 1967 (DAO) (determined by G. Argus). 

This willow was not included in the Vascular Plants of 
Continental Northwest Territories (Porsild and Cody 1980). 
It was however described and mapped by Argus in his 
Guide to the identification of willows in Alaska, the Yukon 
Territory and adjacent regions (Argus 2001). The specimens 
cited above are the basis for the maps in this treatment. 


Salix sphenophylla A.K. Skvort. — MACKENZIE: delta 
low centre polygons, south of Big Lake, 69.3652°N 
134.899°W, D. Bush 25, 2003 (DAO) (determined by 
G. Argus). 

This species was reported as new to the Northwest Terri- 
tories by McJannet et al. (1995) on the basis of a specimen 
collected by E. Kutke at Nig, 74 miles north of Inuvik 
(DAO). The specimen cited above is the second record for 
the Territory. 


POLYGONACEAE 
Polygonum viviparum L., Bistort — KEEWATIN: 
62°08'S0"N 97°53'00"W, K. Reading 37, 15 July 1984 
(DAO); 62°12'00"N 97°52'00"W, K. Reading 51, 3 
July 1984 (DAO). 

The specimens cited above are from sites in the southern 
District of Keewatin about 150 kilometers east of a site just 
east of longitude 100°W (Porsild and Cody 1980). 


CHENOPODIACEAE 

Chenopodium dessicatum A. Nels. (C. leptophyllum 
Nutt.), Narrow-leaved Goosefoot — MACKENZIE: middle of 
open, revegetated sandy slope, 61.7441°N 121.1543°W, 
L. Kershaw 5, 2003. 

Porsild and Cody (1980) reported C. leptophyllum as “In 
the Northwest Territories thus far known only from a Rich- 
ardson specimen collected a Century and a half ago, at Fort 
Franklin, where it was surely an ephemeral introduction’. The 
specimen cited above is from a site about half way between 
Fort Franklin and the west end of Lake Athabasca and may 
also be an ephemeral introduction. 


284 


CARYOPHYLLACEAE 
Cerastium alpinum L. — MACKENZIE: siksik mound, 
Coppermine River Area, 67°32'30"N_ 116°01'30"W, 
K. Reading 77, 29 Aug. 2002 (DAO); Hope Bay Area, 
northeast of Bathurst Inlet, Roberts Lake, 68°10'30"N 
106°33'00"W, K. Reading 241, 7 Aug. 1987 (DAO). 
The first specimen cited above is an extension of the known 
range in the Territory of about 250 kilometers west of a site 
at the south end of Bathurst Inlet. The second specimen is 
an extension of the known range about 170 kilometers to 
the northeast (Porsild and Cody 1980). 


Cerastium beeringianum Cham. & Schlecht. — MAc- 
KENZIE: Hope Bay Area, northeast of Bathurst Inlet, 
Roberts Lake, 68°10'30"N 106°33'00"W, K. Reading 
214, 10 Aug. 1987 (DAO). 

The specimen cited above is an extension of the known 


range in the Territory of about 170 kilometers to the northeast 
of the south end of Bathurst Inlet (Porsild and Cody 1980). 


Minuartia biflora (L.) Schinzl. & Thell. — KEEWATIN: 

11 miles south of east end of Bissett Lake, 63°39'10"N 

95°10'00"W, K. Reading 370, 18 Aug. 1982 (DAO). 
The specimen cited above is an extension of the known 


range in the Territory of about 100 kilometers north of a site in 
the southern half of the Territory (Porsild and Cody 1980). 


Minuartia rossii (R.Br.) Graebn. — KEEWATIN: Bissett 
Creek, 63°46'00"N 95°30'00"W, K. Reading 368, 25 
July 1982 (DAO). 

The specimen cited above is a westerly extension into 
central mainland Keewatin of 500 kilometers from sites on 
Southampton Island. To the north it is known from sites 
adjacent to the Arctic Coast and the Arctic Islands (Porsild 
and Cody 1980). 


Minuartia rubella (Wahlenb.) Graebn. — KEEWATIN: 
south of east end of Bissett Lake, 64°40'30"N 
95°12'00"W, K. Reading 371, 20 Aug. 1982 (DAO). 

The specimen cited above is midway between a site adja- 
cent to the Hudson Bay Coast and about 65°N 98°W (Porsild 
and Cody 1980). 


Minuartia yukonensis Hultén — MACKENZIE: east 
bank of Mackenzie River, 68°52.1'N 134°29.8'W, 
J.V. Matthews s.n., 4 Aug. 1983 (DAO); Mackenzie 
River Valley pipeline survey, Parson’s Lake area, 
68°59'05"N_ 133°30'59"W, J. Lancaster s.n., 5 Aug. 
2002 (DAO); Ya Ya esker, 69.2093°N 134.5636°W, J. 
Lancaster 12A, 9 July 2003 (DAO). 

Porsild and Cody (1980) knew this Amphi-Beringian 
species in the Continental Northwest Territories only as far 
east as the east slope of the Richardson Mountains west of 
the Mackenzie Delta. The specimens cited above extend the 
known distribution eastward to part of the Mackenzie River 
and eastward further to Parson’s Lake about 100 kilometers 
to the northeast. 


Sagina caespitosa (J. Vahl) Lge. — KEEWATIN: TK 
Camp peninsula, Bissett Lake, 63°47'00"N 95°26'30"W, 
K. Reading 367, 17 July 1982 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 125 kilometers east of the 
northernmost site mapped by Porsild and Cody (1980). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Silene walensis (Rupr.) Bouquet (Melandrium apetalum 
(L.) Fenzl ssp. arcticum (Fr.) Hultén) — MACKENZIE: 
Hope Bay Area, northeast of Bathurst Inlet, Roberts 
Lake, 68°10'30"N 106°33'00"W, K. Reading 267, 30 
July 1987 (DAO); KEEwaTIN: wet tundra rills, north of 
30-mile lake, 63°46'00"N 96°07'00"W, K. Reading 
351, 31 July 1981 (DAO). 

The first specimen cited above is an extension of the 
known range in the Territory of about 170 kilometers northeast 
of a site at the south end of Bathurst Inlet (Porsild and Cody 
1980). The Keewatin site is midway between the Hudson 
Bay Coast and sites just east of longitude 100°W. 


Stellaria calycantha (Ledeb.) Bong. — KEEWATIN: Grif- 
fin Lake, 61°19'N 98°46'W, K. Reading 340, 10 Aug. 
1990 (DAO); 62°11'N 97°52'W, K. Reading 61, 19 
July 1984 (DAO); SY area, 62°12'10"N 97°53'00"W, 
K. Reading 136, 5 Aug. 1984 (DAO). 

The specimens cited above extend the known range in 
the Territory about 180 kilometers north of a site north of 
the Manitoba border (Porsild and Cody 1980). 


Stellaria longipes Goldie s.1. — MACKENZIE: scarp foot, 
Coppermine River Area, 67°07'N 115°05'W, K. Read- 
ing 89, 28 Aug. 2002 (DAO); KEEWATIN: east of Kazan 
River, north of Big Bend, 63°45'N 95°38'W, K. Read- 
ing 399B, 8 Aug. 1983 (DAO); south of Whale Lake, 
63°46'N 96°06'W, K. Reading 369, 21 Aug. 1982 
(DAO). 

The first specimen cited above extends the known distri- 
bution about 250 kilometers west of the Burnside Landing 
area adjacent to Bathurst Inlet (Cody et al. 1984). The Kee- 
watin specimens extend the known range to the northeast 
about 250 kilometers to the central area of Keewatin (Porsild 
and Cody 1980). 


NYMPHAEACEAE 

Nymphaea tetragona Georgi ssp. leibergii (Morong) 
Porsild - MACKENZIE: small lake in Camsell Ferry 
infrastructure midway on west shore of lake behind 
beaverlodge, 62.1567°N 122.4908°W, L. Kershaw 
15, 2003 (DAO). 

- This rare plant in the Territory was first reported by A. E. 
Porsild (1939) on the basis of a collection by E. A. Preble 
from an island of the Simpson group 40 miles northeast of 
Resolution, Great Slave Lake. The specimen cited above is 
from a site about 425 kilometers to the west. 


RANUNCULACEAE 
Anemone parviflora Michx. — MACKENZIE: Hope Bay 
Area, northeast of Bathurst Inlet, Windy Lake, 
68°03'15"N 106°37'00"W, K. Reading 296, 14 July 
1987 (DAO); KEEwaTIN: Big Bird Lake, 62°18'00"N 
97°37'40"W, K. Reading 440, 28 June 1985 (DAO). 
The first specimen cited above is an extension of the known 
range in the Territory of about 100 kilometers northeast of a 
site adjacent to Bathurst Inlet (Porsild and Cody 1980). The 
second specimen is an eastward extension of about 80 kilo- 
meters from a site just east of longitude 100°W. 


Anemone richardsonii Hook. — KEEWATIN: creek into 
southwest corner of Pebble Beach Lake, 62°21'50"N 
97°22'00"W, K. Reading 443, July 1985 (DAO). 


2005 


The specimen cited above is encircled by six other sites 
in the southern half of the Territory (Porsild and Cody 1980). 


Caltha natans Pall., Floating Marsh-marigold — KEE- 
WATIN: northwest corner of Bissett Lake, 63°48'00"N 
95°27'00"W, K. Reading 345, 8 Aug. 1983 (DAO). 
This species is rare in the Continental District of Keewatin. 
The specimen cited above is an extension of the known range 
in the Territory of about 375 kilometers to the north and north- 
east from two sites in the south (Porsild and Cody 1980). 


Caltha palustris L. var. arctica (R.Br.) Hultén, Marsh- 
marigold — MACKENZIE: Hope Bay Area, northeast of 
Bathurst Inlet, Roberts Lake, 68°10'30"N_ 106°33'00"W, 
K. Reading 273, 24 July 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 175 kilometers from the south 
end of Bathurst Inlet (Porsild and Cody 1980). 


Ranunculus aquatilis L. var. eradicatus Laest., White 
Water Buttercup — MACKENZIE: lake bottom, Copper- 
mine River Area, 67°32'50"N 116°13'00"W, K. Read- 
ing 75, 23 Aug. 2002 (DAO). 

The specimen cited above is from a site midway between 
the south end of Bathurst Inlet and the east end of Great 
Bear Lake (Porsild and Cody 1980). 


Ranunculus sabinei R.Br. — KEEWATIN: Fly Camp 3, 
65°53'N 98°34'W, S. Smith 204, 20 July 1983 (DAO); 
big esker south of Bissett Lake, 63°40'40"N 95°32'00"W, 
K. Reading 373, 16 Aug. 1981 (DAO); 62°11'00"N 
97°55'00"W, K. Reading 60, 13 July 1984 (DAO). 
The specimens cited above extend the known distribution 


in the Territory south of the Arctic Coast into central District 
of Keewatin (Porsild and Cody 1980). 


Ranunculus xspitzbergensis (Nath.) Hadac — KEEWATIN: 
This hybrid of Ranunculus lapponicus x R. pallasii 
was reported by Cody et al. (1988) on the basis of a 
specimen collected by K. Reading at Jaeger Creek, 
62°11'20"N 97°53'00"W in 1984 and also from north- 
ern Quebec. 


PAPAVERACEAE 
Papaver radicatum Rottb. — MACKENZIE: Hope Bay 
Area, northeast of Bathurst Inlet, Roberts Lake, 
68°10'30"N 106°33'00"W, K. Reading 251, 4 Aug. 
1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory to the northeast of a site about 170 


kilometers from the south end of Bathurst Inlet (Porsild and 
Cody 1980). 


BRASSICACEAE (CRUCIFERAE) 

Braya glabella Richardson — MACKENZIE: Hope Bay 
Area, northeast of Bathurst Inlet, Roberts Lake, 
68°10'30"N 106°33'00"W, K. Reading 485, 12 Aug. 
1987 (DAO) (determined by G.A. Mulligan). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers northeast of the 
south end of Bathurst Inlet (Porsild and Cody 1980) and is 
the northeasternmost site yet known in the Territory. 


CopY AND READING: VASCULAR FLORA OF NWT AND NUNAVUT 


285 


Braya humilis (C.A. Mey.) Robins. — MACKENZIE: 
Hope Bay Area, northeast of Bathurst Inlet, Roberts 
Lake, 68°10'30"N 106°33'00"W, K. Reading 486, 13 
Aug. 1987 (DAO) (determined by G. A. Mulligan). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers northeast of a 
site west of Bathurst Inlet and is the northeasternmost yet 
known in the Territory (Porsild and Cody 1980). To the north 
it is known on Victoria Island. 


Cardamine bellidifolia L. - MACKENZIE: dolomite ridge, 
Coppermine River area, 67°07'00"N 115°45'00"W, 
K. Reading 91, 28 Aug. 2002 (DAO); KEEWATIN: 
62°10'30"N 97°59'00"W, K. Reading 474, 17 Aug. 
1984 (DAO); 62°12'00"N 97°53'00"W, K. Reading 
475, 4 Aug. 1984 (DAO) (determined by G. A. Mulli- 
gan). 

The Mackenzie specimen cited above is an extension of 
the known range in the Territory of about 250 kilometers 
northwest of the south end of Bathurst Inlet. To the west it 
is unknown between there and the Richardson and Mackenzie 
mountains (Porsild and Cody 1980). The Keewatin speci- 
mens cited above are extensions of the known range in the 
Territory of about 70 kilometers east of the southwestern- 
most site mapped by Porsild and Cody (1980). 


Cardamine pratensis L. Lady’s-smock — KEEWATIN: 
62°10'00"N 97°54'00"W, K. Reading 484, 17 July 1984 
(DAO); Pebble Beach Lake, 62°21'58"N 97°30'02"W, 
K. Reading 483, 10 July 1985 (DAO); south of Bis- 
sett Lake, 63°43'N 95°18'W, K. Reading 482, 22 July 
1982 (DAO) (determined by G. A. Mulligan). 

The first two specimens cited above are from sites mid- 
way between sites near the Hudson Bay Coast and longi- 
tude 100°W. The third specimen cited above is the northern- 
most yet found in the Territory from south of the Arctic 
Circle (Porsild and Cody 1980). 


Descurainia sophioides (Fisch. ex Hook.) O.E. Schulz 
— MACKENZIE: Hope Bay Area, northeast of Bathurst 
Inlet, Roberts Lake, 68°10'30"N_ 106°33'00"W, K. 
Reading 254, 3 Aug. 1987 (DAO) (determined by G. 
A. Mulligan). 

The specimen cited above is an extension of the known 
range in the Territory of about 100 kilometers northeast of a 
site on the west side of Bathurst Inlet (Porsild and Cody 1980). 


Draba fladnizensis Wulf — KEEWATIN: 62°12'30"N 
97°52'00"W, K. Reading 507, 3 July 1984 (DAO) 
(determined by G.A. Mulligan). 

The specimen cited above is an extension of the known 
range in the Territory of about 200 kilometers southeast of a 
site adjacent to Chesterfield Inlet (Porsild and Cody 1980) 
(determined by G. A. Mulligan). 


Draba juvenalis Kom. (D. longipes Raup) — MACKEN- 
ZiE: Hope Bay Area, northeast of Bathurst Inlet, Roberts 
Lake, 68°10'30"N 106°33'00"W, K. Reading 500, 12 
Aug. 1987 (DAO) (determined by G. A. Mulligan). 

The specimen cited above is an extension of the known 
range in the Territory of about 500 kilometers northeast of 
the east end of Great Bear Lake. To the northeast it is known 
from two localities on southeastern Victoria Island (Porsild 
and Cody 1980). 


286 


Draba nivalis Liljebl. — KEEWATIN: Pebble Beach 
Lake, 62°21'40"N 97°29'54"W, K. Reading 511, 28 
June 1985 (DAO); 62°11'00"N 97°55'00"W, K. Read- 
ing 510, 13 July 1984 (DAO) (determined by G. A. 
Mulligan). 

The specimens cited above are from sites midway 
between sites in the central area of the Territory south of the 


Arctic Circle and a site in the southeast adjacent to latitude 
100°W (Porsild and Cody 1980). 


Draba wahlenbergii Hartm. (D. lactea Adams) — MAc- 
KENZIE: Hope Bay Area, northeast of Bathurst Inlet, 
Roberts Lake, 68°10'30"N 106°33'00"W, K. Reading 
497, 13 July 1990 (DAO); K. Reading 501, 13 Aug. 
1987 (DAO); K. Reading 508, 12 Aug. 1987 (DAO); 
KEEWATIN: Camp 3, 61°12'N 97°57'W, K. Reading 
499, 13 July 1990 (DAO); 62°10'30"N 97°59'00"W, 
K. Reading 502, 17 Aug. 1984 (DAO); Imikula Lake, 
62°14'00"N 97°37'00"W, K. Reading 504, 18 June 
1985 (DAO); Big Bird Lake, 62°17'30"N 97°39'00"W, 
K. Reading 503, 27 June 1985 (DAO); Pebble Beach 
Lake, 62°22'00"N 97°30'00"W, K. Reading 506, 7 
July 1985 (DAO) (determined by G. A. Mulligan). 

The District of Mackenzie specimens cited above are from 
sites about 170 kilometers northeast of the south end of 
Bathurst Inlet. To the north it is known from southern Victoria 
Island and is widespread throughout the Arctic Islands. The 
District of Keewatin specimens are sites midway between 
sites adjacent to the Hudson Bay Coast and the Mackenzie- 
Keewatin border and south of sites north of latitude 63°N 
(Porsild and Cody 1980). 


Erysimum pallasii (Pursh) Fern. - MACKENZIE: Hope 
Bay Area, northeast of Bathurst Inlet, Windy Lake, 
68°03'11"N 106°37'00"W, K. Reading 298, 14 July 
1987 (DAO); KEEwaATIN: north side of Mire Creek, 
61°21'00"N 97°53'30'W, K. Reading 514, 2 July 1990 
(DAO) (determined by G. A. Mulligan). 

The first specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers northeast of 
the south end of Bathurst Inlet (Porsild and Cody 1980). The 
second specimen is a new record for the District of Keewatin 
and is an extension of about 765 kilometers southeast of the 
south end of Bathurst Inlet. 


Eutrema edwardsii R.Br. — MACKENZIE: Hope Bay 
Area, northeast of Bathurst Inlet, Windy Lake, 
68°03'15"N 106°37'00"W, K. Reading 295, 14 Aug. 
1987 (DAO) (determined by G. A. Mulligan). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers to the north- 
east from the south end of Bathurst Inlet. To the north it is 
known on Victoria Island and it is widespread on the Arctic 
Islands (Porsild and Cody 1980). 


Parrya arctica R.Br. — MACKENZIE: Hope Bay Area, 
northeast of Bathurst Inlet, Roberts Lake, 68°10'30"N 
106°33'00"W, K. Reading 476, 10 Aug. 1987 (DAO) 
(determined by G. A. Mulligan). 

The specimen cited above is an extension of the known 
range in the Territory of about 375 kilometers east of a site 
west of Coronation Gulf. To the north it is known from Vic- 
toria Island and is widespread on the Arctic Islands west of 
longitude 90°W (Porsild and Cody 1980). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


SAXIFRAGACEAE 

Chrysosplenium tetrandrum (Lund) Fries — MACKENZIE: 

rocky lakeshore, Coppermine River Area, 67°33'30"N 

116°45'00"W, K. Reading 44, 21 Aug. 2002 (DAO). 
The specimen cited above is from a site midway between 

the south end of Bathurst Inlet and Coppermine (Porsild and 

Cody 1980). 


Saxifraga nelsoniana D. Don ssp. _ porsildiana 
(Calder and Savile) Hultén (S. punctata L. ssp. por- 
sildiana Calder & Savile) — KEEWATIN: Pebble Beach 
Lake, south of Yathkyed Lake, 62°22'00"N 97°30'00"W, 
K. Reading 175, 22 July 1985 (DAO). 

The specimen cited above is an extension of about 250 
kilometers south of the site reported by Cody et al. (2003). 


Saxifraga nivalis L., Alpine Saxifrage — MACKENZIE: 
wet tundra below scarp, Coppermine River Area, 
67°31'12"N 116°09'58"W, K. Reading 64, 22 Aug. 
2002 (DAO); scarp foot wet, Coppermine River Area, 
67°32'28"N 116°01'00"W, K. Reading 82, 30 Aug. 
2002 (DAO). 

The specimens cited above are from sites midway between 
Bathurst Inlet and Coppermine (Porsild and Cody 1980). 


Saxifraga rivularis L. — KEEWATIN: wet shaded rocky 
ravine south of Wolf Lake, 63°39'00"N 95°11'00"W, 


K. Reading 360, 16 Aug. 1982 (DAO); big esker south | 


of Bissett Lake, 63°40'40"N 95°32'00"W, K. Reading 
374, 16 Aug. 1981 (DAO); east of Bissett Lake, | 
63°46'00"N 95°46'00"W, K. Reading 342, 27 July 
1982 (DAO). 

The specimens cited above are from sites midway between 


the Hudson Bay Coast and sites adjacent to longitude 100°W 
(Porsild and Cody 1980). 


ROSACEAE 
Potentilla biflora Willd. — MACKENZIE: wet tundra, — 
Coppermine River Area, 66°43'10"N 114°58'00"W, © 
K. Reading 523, 7 Sept. 2002 (DAO); rocky tundra and — 
basalt outcrops, Coppermine River Area, 67°21'00"N_ | 
115°51'00"W, K. Reading 2, 17 Aug. 2002 (DAO); | 
scarp foot scree, 67°30'10"N 116°10'00"W, K. Reading | 
524, 22 Aug. 2002 (DAO); clay tundra boils, Copper- | 
mine River Area, 67°31'30"N 116°14'10"W, K. Read- | 
ing 525, 19 Aug. 2002 (DAO); Dismal Lakes, 67°29'N_ | 
117°36'W, L.W. Stock s.n., 8 Aug. 1947 (DAO). 

The nearest sites of this species to those listed above in | 
this Territory are in the vicinity of the Bluenose Lake Area ! 
about 500 kilometers to the northwest (Cody et al. 2003). i 
There is also a site to the northeast on Victoria Island (Por- | 
sild and Cody 1980). 


Potentilla nivea L. ssp. nivea — KEEWATIN: high dry | 
esker-top, east of Kazan Falls, 63°41'N 95°38'W, K. | 
Reading 354, 30 Aug. 1982 (DAO); west southwest of 
Bissett Lake, 63°44'30"N 95°28'00'W, K. Reading 334, 
4 Aug. 1982 (DAO); TK camp peninsula, Bissett Lake, 
63°47'00"N 95°26'30"W, K. Reading 379, 9 Aug. 1982 | 
(DAO). 

The specimens cited above are an extension of the known | 
range in the Territory of about 170 kilometers northeast of a 
site mapped by Porsild and Cody (1980). 


2005 


Potentilla palustris (L.) Scop., Marsh Fivefinger — 
KEEWATIN: south of Deep Bay of Kazan River, 
63°41'30"N 95°37'00"W, K. Reading 333, 3 Aug. 
1982 (DAO). 

The specimen cited above is an extension of the known 
range of about 170 kilometers northeast of a site in central 
Keewatin mapped by Porsild and Cody (1980). 


Rosa blanda Ait. — MACKENZIE: Mackenzie River Valley 
survey, Creek bank, 64°35'23"N 124°48'22"W, B. Cor- 
nish s.n., 17 July 2002 (DAO). 

Porsild and Cody (1980) knew this species in the Territory 
only from the vicinity of Wrigley. The specimen cited above 
is an extension of the known range of about 160 kilometers 
to the northwest. 


Rubus arcticus L. ssp. acaulis (Michx.) Focke — KEE- 
WATIN: Griffin Lake, 61°14'55"N 98°46'00"'W, K. Read- 
ing 89, 10 Aug. 1990 (DAO). 

The specimen cited above is from a site midway between 
two sites mapped by Porsild and Cody (1980) in southwest- 
ern District of Keewatin. 


Rubus idaeus L. s.l. (R. strigosus Michx.), Wild Red 
Raspberry — KEEWATIN: Griffin Lake, 61°18'40"N 
98°52'00"W, K. Reading 532, 14 Aug. 1990 (DAO). 
Porsild and Cody (1980) knew this species from a single 
location in the southwest of the Territory. The specimen cited 
above is an extension of about 100 kilometers to the north. 


FABACEAE (LEGUMINOSAE) 

Astragalus australis (L.) Lam. (A. richardsonii Shel- 
don) — MACKENZIE: Hope Bay Area, northeast of Bath- 
urst Inlet, Roberts Lake, 68°10'30"N 106°33'00"W, 
K. Reading 264, 30 July 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Porsild and Cody 1980) of about 170 
kilometers northwest of the south end of Bathurst Inlet. To 
the north it is also known from Victoria Island. 


Astragalus eucosmus Robins. — KEEWATIN: west side 
of Kneeling Man Lake, south of Yathkyed Lake, 
62°19'30"N 97°33'20"W, K. Reading 187A, 17 July 
1985 (DAO); “No-camp” Lake south of Yathkyed 
Lake, K. Reading 85-17, 15 July 1985 (DAO); “‘Peb- 
ble Beach” Lake, south of Yathkyed Lake, 62°22'M 
97°30'W, K. Reading 85-26, 17 July 1985 (DAO); 
vicinity of Meliadine Lake, 63°01'44"N 92°10'5 1" W, 
K. Reading 94-26, 1984 (DAO); on rock outcrop, Ran- 
kin Inlet 5 km northwest of town, 62°49'N 92°05'W, 
J. B. Korol s.n., 19 July 1988 (DAO). 

The specimens cited above are the first known from the 
Continental District of Keewatin (Porsild and Cody 1980). 


| Astragalus eucosmus Robins. f. albinus Fern. — KEE- 
| WATIN: north end of No-camp Lake, south of Yathkyed 
| Lake, 62°22'15"N 97°27'10"W, K. Reading 382, 15 
July 1985 (DAO). 

This white-flowered form was described by Fernald (1926) 
on the basis of a specimen from Ha-Ha Mountain in New- 
foundland. The specimen cited above is the first known from 
| the Continental Northwest Territories. 


Hedysarum alpinum L., Liquorice-root — MACKENZIE: 
Hope Bay Area, northeast of Bathurst Inlet, Roberts 


CoDY AND READING: VASCULAR FLORA OF NWT AND NUNAVUT 


287 


Lake, 68°10'30"N_ 106°33'00"W, K. Reading 236, 8 
Aug. 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 80 kilometers from a site in 
the middle of Bathurst Inlet. It is the most northeastern site 
yet known in the Territory. 


Hedysarum boreale Nutt. ssp. mackenzii (Richards.) 
Welsh (H. mackenzii Richards.) — MACKENZIE: Hope 
Bay Area, northeast of Bathurst Inlet, Roberts Lake, 
68°10'30"N 106°33'00"W, K. Reading 237, 8 Aug. 
1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers northeast of 
the south end of Bathurst Inlet (Porsild & Cody 1980). To the 
north it is known on Victoria Island. 


Lupinus arcticus Wats. — MACKENZIE: Hope Bay 
Area, northeast of Bathurst Inlet, Roberts Lake, 
68°10'30"N 106°33'00"W, K. Reading 237, 2 Aug. 
1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers to the north- 
east of the south end of Bathurst Inlet. To the northeast it is 
known on Victoria Island (Porsild and Cody 1980). 


Oxytropis arctobia Bunge — MACKENZIE: George Lake, 
south of Bathurst Inlet at the head of Western River, 
approx. 65°57'20"N_ 107°30'W, K. Reading 451, 18 
Aug. 1988 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 75 kilometers south of the 
south end of Bathurst Inlet (Porsild and Cody 1980). 


Oxytropis deflexa (Pall.) DC. ssp. foliosa (Hook.) Cody 
— MACKENZIE: Amphibolite ridge crown, north of the 
east arm of Great Slave Lake, 62°42'N 113°05'W, K. 
Reading 12, 13 Sept. 2002 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 27 kilometers east of a site in 
the vicinity of Yellowknife (Cody et al. 2003). 


Oxytropis maydelliana Trautv. — MACKENZIE: Hope 
Bay Area, northeast of Bathurst Inlet, Roberts Lake, 
68°10'30"N 106°33'00"W, K. Reading 224, 8 Aug. 
1987, 244, 6 Aug. 1987 (DAO). 

The specimens cited above are an extension of the known 
range in the Territory of about 170 kilometers northeast of 
the south end of Bathurst Inlet. To the northeast it is known 
on Victoria Island (Porsild and Cody 1980). 


CALLITRICHACEAE 
Callitriche hermaphroditica L. — MACKENZIE: silty 
lake bottom, Coppermine River Area, 67°32'55"N 
116°13'00"W, K. Reading 540, 23 Aug. 2002 (DAO). 
The specimen cited above which is the northwesternmost 
yet found in the Territory, is an extension of the known 
range of about 170 kilometers to the northwest from a site 
near the east end of Great Bear Lake (Porsild and Cody 1980). 


ELEAGNACEAE 

Shepherdia canadensis (L.) Nutt., Soapberry — KEE- 
WATIN: Camp 2, 61°21'00"N 97°53'30"W, K. Reading 
542, 4 July 1990 (DAO). 


288 


The species is new to the flora of mainland Keewatin. It 
is a southeastward extension of the known range of about 425 
kilometers from a site near the eastern limit of the District 
of Mackenzie (Porsild and Cody 1980). 


ONAGRACEAE 
Epilobium davuricum Fisch. — KEEWATIN: dark frost- 
boils, T-Bone Lake, SY, 62°13'N 97°53'W, K. Read- 
ing 390, 15 Aug. 1984 (DAO). 

The specimen cited above is from a site midway between 
two sites in the Territory reported by Cody et al. (2003). 


Epilobium latifolium L. f. albiflorum Nath. — KEE- 
WATIN: Griffin Lake, 61°14'30"N 98°38'00"W, K. 
Reading 544, 17 Aug. 1990 (DAO). 

Porsild and Cody (1980) stated that the flowers of this 
species were purple or rarely white, but no localities of the 
white flower form have previously been reported for this 
region. 


Epilobium palustre L., Swamp Willowherb — MACKEN- 
ZIE: Hope Bay Area, northeast of Bathurst Inlet, 
Roberts Lake, 68°10'30"N 106°33'00"W, K. Reading 
247, 6 Aug. 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers to the north- 
east of a site near the south end of Bathurst Inlet (Porsild 
and Cody 1980). 


HALORAGACEAE 

Myriophyllum alterniflorum DC. — MACKENZIE: Pan- 
handle Lake, SE of Parsons, 68.9104°N 121.555°W, 
J. Lancaster 13, 2003 (DAO). 

This is a rare species in the Territory (McJannet et al. 1995). 
The specimen cited above is from a site about 185 kilome- 
ters northwest of a site adjacent to the northeast end of 
Great Bear Lake. 


PYROLACEAE 

Moneses uniflora (L.) Gray, One-flowered Pyrola — 
KEEWATIN: Griffin Lake, 61°16'40"N 98°42'00"W, K. 
Reading 551, 22 July 1990 (DAO); in white spruce 
copses on tundra, very local, small colonies, north of 
Griffin Lake, 61°02'N 98°12'W, K. Reading s.n., 15 
Aug. 1990 (DAO). 

This species is new to Continental District of Keewatin. 
The nearest sites known to Porsild and Cody (1980) were 
near the east end of Great Slave Lake about 700 kilometers to 
the northwest and adjacent to Hudson Bay in the vicinity of 
Churchill, Manitoba, about 250 kilometers to the southeast. 


Pyrola secunda L. s.1. — KEEWATIN: north of Bissett 
Lake, 63°48'30"N 95°24'00"W, K. Reading 364, 13 
Aug. 1982 (DAO). 

The specimen cited above is from the central part of the 
District of Keewatin about 200 kilometers south and 250 kilo- 
meters northeast of sites mapped by Porsild and Cody (1980). 


ERICACEAE 

Arctostaphylos alpina (L.) Spreng., Alpine Bearberry 
— MACKENZIE: Hope Bay Area, northeast of Bathurst 
Inlet, Windy Lake, 68°03'15"N 106°37'00"W, K. Read- 
ing 291, 7 July 1987 (DAO). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


The specimen cited above is an extension of the known 
range in the Territory of about 80 kilometers to the north- 
east of a site adjacent to Bathurst Inlet (Porsild and Cody 
1980). 


Ledum groenlandicum Oeder, Labrador-tea — KEE- 
WATIN: UG Lake, 62°45'00"N 98°40'00"W, K. Read- 
ing 454, date unknown (DAO); lakeshore, northwest 
of Imikula Lake, 62°02'N 97°40'W, K. Reading 459, 
July-August 1984 (DAO). 

The specimens cited above are the northernmost yet found 
in the Territory (Porsild and Cody 1980). 


Vaccinium vitis-idaea L. var. minus Lodd., Mountain 
Cranberry — MACKENZIE: Hope Bay Area, northeast of 
Bathurst Inlet, Roberts Lake, 68°10'30"N 106°33'00"W, 
K. Reading 282, 22 July 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 80 kilometers to the east of a 
site on the west side of Bathurst Inlet (Porsild and Cody 1980). 


GENTIANACEAE 

Gentiana affinis Griseb. — MACKENZIE: Mackenzie 
River Valley pipeline survey, stream bed and low ter- 
races along small creek/with gravel bars upstream, 
63°22'38"N 123°29'35"W, L. Kershaw s.n., 6 Aug. 2002 
(DAO); riparian fluvial silt along river, 66.7082°N 
129.3002°W, L. Kershaw 8, 2003 (DAO). 

McJannet et al. (1995) knew this rare species in the 
Northwest Territories from locations at the west end of Great 
Slave Lake in the vicinity of Fort Good Hope, Mackenzie 
Mountains in the vicinity of the Keele River and near Fort 
Norman. The first specimen cited above is from north of 
Wrigley. The second specimen is the northernmost yet found 
in the Territory and is an extension of the known range of 
about 150 kilometers northwest of Fort Good Hope. 


LAMIACEAE 
Lycopus uniflorus Michx., Tuberous Water-horehound - 
— MACKENZIE: muddy run south side of beaver lodge » 
near NW corner lake, 62.9858°N 123.1755°W, L. 
Kershaw 11, 2003 (DAO). 

This is a rare species in the Territory (McJannet et al. 
1995), previously known from only east of the Slave River 
(Porsild and Cody 1980). The specimen cited above is an 
extension of the known range in the Territory of about 635 
kilometers to the northwest. 


SCROPHULARIACEAE 
Lagotis glauca Gaertn. ssp. minor (Willd.) Hultén — | 
MACKENZIE: hill west of Canoe Lake, head of creek, 
68°15'N 135°54'W, V. J. Krajina s.n., 11 Aug. 1965 
(DAO). 

The specimen cited above is from a site about 100 kilome- 
ters north of a site mapped by Porsild & Cody (1980). This | 
site is the northernmost yet found in the Territory. 


Limosella aquatica L., Mudwort — MACKENZIE: black | 
spruce/Labrador Tea/cloudberry area in strip of ex- 
posed mud, 63.4867°N 123.6914°W, L. Kershaw 10, 
2003 (DAO). 

This is a rare species in the Territory (McJannet et al. 1995). 
The specimen cited above is from a site midway between the | 


2005 


east end of Great Bear Lake and a site adjacent to the Canol 
Road in the Yukon Territory. 


Pedicularis capitata Adams — MACKENZIE: Hope Bay 
Area, northeast of Bathurst Inlet, Roberts Lake, 
68°10'30"N 106°33'00"W, K. Reading 271, 28 July 
1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 80 kilometers east of a site 
on the west side of the north end of Bathurst Inlet (Porsild 
and Cody 1980). 


Pedicularis langsdorfii Fisch. ssp. arctica (R.Br.) Pen- 
nell ex Hultén (P. arctica R.Br.) — MACKENZIE: Hope 
Bay Area, northeast of Bathurst Inlet, Roberts Lake, 
68°10'30"N 106°33'00"W, K. Reading 272, 26 July 
1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers northeast of a 
site at the south end of Bathurst Inlet (Porsild and Cody 1980). 


Pedicularis lapponica L., Lapland Lousewort — MAc- 
KENZIE: Hope Bay Area, northeast of Bathurst Inlet, 
Roberts Lake, 68°10'30"N 106°30'00"W, K. Reading 
234, 8 Aug. 1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers northeast of 
the south end of Bathurst Inlet (Porsild and Cody 1980). 


Pedicularis macrodonta Richards. (P. parviflora Smith ex 
Rees var. macrodonta (Richards.) Welsh — MACKENZIE: 
in graminoid/shrub fen with Juncus stygius, 60.6382°N 
120.34633°W, D. Soppet 16, 17 Aug. 2003 (DAO). 

This is a rare species in the District of Mackenzie (McJan- 
net et al. 1995). The specimen cited above is only the second 
known in the Territory and is about 200 kilometers west of 
the west end of Great Slave Lake. 


RUBIACEAE 

~Galium trifidum L. — KEEWATIN: among shoreline rocks 

just north of TK camp, Bissett Lake, 63°47'02"N 

95°26'31"W, K. Reading 358, 30 Sept. 1982 (DAO). 
The specimen cited above is the northernmost yet found in 

the Territory (Porsild and Cody 1980). It is about 300 kilo- 

meters north of the only site known to Porsild and Cody 

(1980). 


CAPRIFOLIACEAE 

Linnaea borealis L. var. americana (Forbes) Rehd., 
Twinflower — MACKENZIE: spruce “island” on dolomite 
ridge, Coppermine River Area, 67°07'00"N 115°45'00"'W, 
| K. Reading 84, 28 Aug. 2002 (DAO). 

The specimen cited above is the most northeasterly yet 
: found in the Territory (Porsild and Cody 1980). It is from a 
| Site about 250 kilometers northeast of the east end of Great 
| Bear Lake. 


| Campanula uniflora L. — KEEWATIN: west southwest 
of Bissett Lake, 63°43'20"N 95°30'00"W, K. Reading 
337, 4 Aug. 1982 (DAO). 

The specimen cited above is from a site midway between 
} 4 Site adjacent to the Hudson Bay Coast and a site adjacent 
to longitude 100°W (Porsild and Cody 1980). 


CODY AND READING: VASCULAR FLORA OF NWT AND NUNAVUT 


289 


ASTERACEAE 
Antennaria isolepis Greene — KEEWATIN: south of east 
end of Bissett Lake, 63°43'00"N 95°11'00"W, K. Readi- 
ng 403, 15 Aug. 1982 (DAO); south of Bissett Lake, 
63°41'30"N 95°18'00"W, K. Reading 418, 22 July 1982 
(DAO); 10 miles northeast of Bissett Lake, 63°56'99"N 
95°07'00" W, K. Reading 405, 3 July 1981 (DAO). 

The specimens cited above are from sites about 200 kilo- 


meters northeast and southeast of sites about longitude 99°W 
in the Territory (Porsild and Cody 1980). 


Arnica angustifolia Vahl in Hornem. ssp. attenuata 
(Greene) Douglas & Ruyle-Douglas (A. alpina (L.) 
Olin ssp. attenuata (Greene) Maguire) — MACKENZIE: 
Coppermine River Area, scarp slope, K. Reading 33, 
19 Aug. 2002 (DAO); foot of scarp in sand, 67°31'10"N 
116°09'54"W, K. Reading 54, 22 Aug. 2002 (DAO); 
sand/gravel ridge, 67°23'45"N 116°27'00"W, K. Read- 
ing 32, 16 Aug. 2002 (DAO). 

The specimens cited above extend the known distribution 
in the Territory about 170 kilometers to the northeast from a 
site adjacent to the southeast end of Great Bear Lake (Por- 
sild and Cody 1980). 


Arnica chamissonis Less. — MACKENZIE: Yellowknife 
Area, roadside clearing, Boulder Bay, 62°32'00"N 
113°20'00"W, K. Reading 11, 11 Sept. 2002 (DAO). 

The specimen cited above is from a site about 170 kilo- 
meters northwest of a site on the south side of eastern Great 
Slave Lake. 


Artemisia furcata Bieb. (A. hyperborea Rydb.) — 
MACKENZIE: Coppermine River Area, 67°32'30"N 
116°00'00"W, K. Reading 582, 29 Aug. 2002 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 160 kilometers to the southeast 
of a site west of Coppermine and northeast of a site at the 
west end of Great Bear Lake (Porsild and Cody 1980). 


Artemisia tilesii Ledeb. s.1. - MACKENZIE: Hope Bay 
Area, northeast of Bathurst Inlet, 68°10'30"N 
106°33'00"W, K. Reading 248, 3 Aug. 1987 (DAO). 
The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers northeast of a 
site near the south end of Bathurst Inlet (Porsild and Cody 
1980). It is the northeasternmost site yet known in the Territory. 


Aster puniceus L., Purple-stemmed Aster — MACKENZIE: 
in calcareous area where stream widens and pools on 
flat area before flowing into mineral lake, Wrigley area, 
62.7307°N 123.1098°W, D. Soppet 26B, 2003 (DAO). 

Porsild and Cody (1980) suggested that this species should 
be looked for on the Salt Plain west of Fort Smith. The spec- 
imen cited above is from a site about 730 kilometers north- 
west of a site mapped near the west end of Lake Athabasca 
by Porsild and Cody. 


Aster pygmaeus Lindl. — MACKENZIE: Hope Bay Area, 
northeast of Bathurst Inlet, Roberts Lake, 68°10'30"N 
106°33'00"W, K. Reading 225, 9 Aug. 1987 (DAO). 
The specimen cited above is an extension of the known 
range in the Territory of about 125 kilometers northeast of a 
site about the middle of Bathurst Inlet and it is the northeast- 
ernmost yet known in the Territory (Porsild and Cody 1980). 


290 


Aster sibiricus L. — MACKENZIE: Coppermine River 
Area, foot of scarp, sandy, 67°31'10"N 116°09'54"W, 
K. Reading 55, 22 Aug. 2002 (DAO); sandy slope near 
scarp, 67°32'50"N 116°13'00"W, K. Reading 584, 23 
Aug. 2002 (DAO). 

The specimens cited above are from sites midway between 


the south end of Bathurst Inlet and the vicinity of Copper- 
mine (Porsild and Cody 1980). 


Crepis nana Richards. — MACKENZIE: Hope Bay Area, 
northeast of Bathurst Inlet, Windy Lake, 68°03'15"N 
106°37'00"W, K. Reading 300, 14 July 1987 (DAO); 
George Lake, south of Bathurst Inlet at the head of 
Western River, 65°57'20"N, 107°30'W, K. Reading 
453, 18 Aug. 1988 (DAO); sandy ridge, Coppermine 
River Area, 67°24'00"N 116°24'00"W, K. Reading 
586, 18 Aug. 2002 (DAO). 

The first specimen cited above is an extension of the 
known range in the Territory of about 170 kilometers north- 
east of the south end of Bathurst Inlet. The second specimen 


is from a site about 170 kilometers northwest of the south 
end of Bathurst Inlet (Porsild and Cody 1980). 


Erigeron uniflorus L. ssp. eriocephalus (Vahl. ex Hor- 
nem.) Crong. (E. eriocephalus Vahl. ex Hornem.) — 
MACKENZIE: Coppermine River Area, sand ridge near 
creek, 67°23'00"N 116°27'00"W, K. Reading 595, 16 
Aug. 2002 (DAO). 

The specimen cited above is from a site midway between 


the vicinity of Coppermine and the south end of Bathurst 
Inlet (Porsild and Cody 1980). 


Matricaria ambigua (Ledeb.) Kryl., Sea-shore Camo- 
mile — MACKENZIE: Hope Bay Area, northeast of Bath- 
urst Inlet, Roberts Lake, 68°10'30"N 106°33'00"W, 
K. Reading 268, 30 July 1987 (DAO); Windy Lake, 
68°03'15"N 106°37'00"W, K. Reading 290, 7 July 1987 
(DAO). 

The specimens cited above are an extension of the known 


range in the Territory of about 170 kilometers northeast of 
the south end of Bathurst Inlet (Porsild and Cody 1980). 


Senecio congestus (R.Br.) DC. — MACKENZIE: Hope Bay 
Area, northeast of Bathurst Inlet, Roberts Lake, 
68°10'30"N 106°33'00"W, K. Reading 232, 10 Aug. 
1987 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 170 kilometers to the northeast 
from the south end of Bathurst Inlet (Porsild and Cody 1980). 


Solidago graminifolia (L.) Salisb. var. major (Michx.) 
Fern. (Euthamis graminifolia (L.) Nutt. var. gramini- 
folia) — MACKENZIE: clearing with coarse rocks as soil 
base, #3 bay south of R.R.T.P. Camp, Tsu Lake, 
60°35'N 111°52'W, J. Thompson s.n., 19 Aug. 1980 
(DAO); rocky shoreline, 250 metres west of Tsu Lake 
camp, 60°35'N 111°52'W, D. Campbell s.n., 19 Aug. 
1980 (DAO); river banks, 63.4719°N 123.7029°W, L. 
Kershaw 27, 2003. 

Porsild and Cody (1980) and McJannet et al. (1995) knew 
this rare plant in the Territory only from the vicinity of Fort 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Simpson. The first two specimens cited above are from sites 
northeast of Lake Athabasca. The third specimen is a north- 
ward extension adjacent to the Mackenzie River of about 
100 kilometers from the northernmost mapped by Porsild 
and Cody (1980). 


Acknowledgments 

We thank the many individuals who have contri- 
buted specimens to the DAO Herbarium; George Argus 
for the identification of Salix; Gerry Mulligan for the 
identification of Cruciferae; Leslie Cody for the many 
hours inputting this information on her computer; and 
Paul Catling for reviewing an earlier version of this 
manuscript. 


Literature Cited 

Argus, G. W. 2001. A Guide to the identification of willows 
in Alaska, the Yukon Territory and adjacent regions (pub- 
lished privately). 

Cody, W. J., M. Blondeau, and J. Cayouette. 1988. Ranun- 
culus x spitzbergensis (Nath.) Hadac, An addition to the 
flora of North America. Rhodora 90: 27-36. 

Cody, W. J., K. L. MacInnes, J. Cayouette, and S. Dar- | 
byshire. 2000. Alien and Invasive Native Plants along © 
the Norman Wells Pipeline, District of Mackenzie, North- 
west Territories, Canadian Field-Naturalist 114: 126-137. 

Cody, W. J., K. L. Reading, and J. M. Line. 2003. Addi- 
tions and Range Extensions to the Vascular Plant Flora of — 
the Continental Northwest Territories and Nunavut, Canada 
II. Canadian Field-Naturalist 117: 448-465. | 

Cody, W. J., G. W. Scotter, and S. C. Zoltai. 1984. Additions — 
to the Vascular Plant Flora of Bathurst Inlet, Northwest 
Territories, Canada. Canadian Field-Naturalist 98: 171-177. 

Cody,.W. J., G. W. Scotter, and S. C. Zoltai. 1992. Vascu- 
lar Plant Flora of the Melville Hills Region, Northwest 
Territories. Canadian Field-Naturalist 106: 87-99. | 

Cody, W. J. and S. S. Talbot. 1978. Vascular Plant Range | 
Extensions to the Heart Lake Area, District of Mackenzie, | 
Northwest Territories. Canadian Field-Naturalist 92: 137- | 
143. 

Haynes, R. R. 1974. A Revision of North American Pota- | 
mogeton subsection Pusilli (Potamogetonaceae). Rhodora | 
76: 564-649. 

Fernald, M. L. 1926. Two summers of botanizing in New- | 
foundland. Rhodora 28: 215. | 

McJannet, C. L., G. W. Argus, and W. J. Cody. 1995. | 
Rare Vascular Plants in the Northwest Territories. Syllo- — 
geus 73, Canadian Museum of Nature, Ottawa. 104 pages. 

Porsild, A. E. 1939. Notes on the occurrence of Zostera and | 
Zanichellia in arctic North America. Rhodora 73: 90-94. | 

Porsild, A. E., and W. J. Cody. 1980. Vascular Plants of | 
Continental Northwest Territories, Canada. National | 
Museum of Natural Sciences, Ottawa. 667 pages. i 

Scotter, G. W., and W. J. Cody. 1974. Vascular Plants of | 
Nahanni National Park and Vicinity, Northwest Territories, | 
Naturaliste Canadien 101: 861-891. ! 

Thieret, J. W. 1961. New Plant Records for Southwestern ; 
District of Mackenzie, Canadian Field-Naturalist 75: 111- — 
121: 


Received 20 October 2004 
Accepted 11 March 2005 


Notes 


Apparent Predation by Gray Jays, Perisoreus canadensis, on 
Long-toed Salamanders, Ambystoma macrodactylum, in the Oregon 
Cascade Range 


MICHAEL P. MurRAY!, CHRISTOPHER A. PEARL”, and R. BRUCE BuRY? 


' Crater Lake National Park, P.O. Box 7, Crater Lake, Oregon 97604 USA (e-mail: Michael_Murray @nps.gov). Author for 
correspondence 
> USGS Forest and Rangeland Ecosystem Science Center, 3200 SW Jefferson Way, Corvallis, Oregon 97331 USA 


Murray, Michael P., Christopher A. Pearl, and R. Bruce Bury. 2005. Apparent predation by Gray Jays, Perisoreus canadensis, 
on Long-toed Salamanders, Ambystoma macrodactylum, in the Oregon Cascade Range. Canadian Field-Naturalist 
1192): 291-292. 


We report observations of Gray Jays (Perisoreus canadensis) appearing to consume larval Long-toed Salamanders (Ambystoma 
macrodactylum) in a drying subalpine pond in Oregon, USA. Corvids are known to prey upon a variety of anuran amphibians, 
but to our knowledge, this is the first report of predation by any corvid on aquatic salamanders. Long-toed Salamanders 
appear palatable to Gray Jays, and may provide a food resource to Gray Jays when salamander larvae are concentrated in 
drying temporary ponds. 


Key Words: Gray Jay, Perisoreus canadensis, Long-toed Salamander, Ambystoma macrodactylum, amphibian, corvid, diet, 


larvae, pond. 


Corvid birds are generalist feeders and “oppor- 
tunistic” predators (Bent 1946; Marshall et al. 2003). In 
montane regions of western North America, corvids 
sometimes consume anuran amphibians, which can 
offer a concentrated food resource in breeding or larval 
rearing aggregations (Beiswenger 1981; Olson 1989; 
Corn 1993). Common Ravens (Corvus corax) and 
American Crows (C. brachyrhynchos) are predators of 
adult Western Toads (Bufo boreas) (Olson 1989; Corn 
1993; Brothers 1994). Clark’s Nutcrackers (Nucifraga 
columbiana) consume larval Columbia Spotted Frogs 
(Rana luteiventris) (Turner 1960; Pilliod 2002). Gray 
Jays (Perisoreus canadensis) prey upon larval B. boreas 
(Beiswenger 1981) and juvenile Boreal Chorus Frogs 
(Pseudacris triseriata) (Tordoff 1980). To our knowl- 
edge, there are no reports of corvids feeding on aquatic 
stages of salamanders. Here, we report Gray Jay pre- 
dation on larval Long-toed Salamanders (Ambystoma 
macrodactylum) in the southern Cascade Range in 
Oregon. 

We observed Gray Jay feeding behavior in the early 
afternoon of 2 October 2003 at a temporary pond on 
Whitehorse Bluff in Crater Lake National Park (ele- 
vation 1933 m; UTM Zone 10, E565248, N4747116; 
NAD27). This pond is located on a terrace with numer- 
ous other temporary ponds within a forest dominated 
by old-growth Mountain Hemlock (Tsuga merten- 
siana). When full after snowmelt, the observation pond 
covers an area of approximately 800 m? and has a 
maximum depth of 0.5 — 1 m. At the time of our obser- 


vations, the pond had dried to a small pool (approxi- 
mately 40 m*) which was about 5% of the area cov- 
ered when full. We observed two Gray Jays foraging 
at a muddy circular depression (0.3 m diameter) 
located 2 m from the only remaining pool in the pond 
basin. This depression was separated from the pool 
by dried mud. We observed no other similar saturated 
depressions within the pond basin. Each jay dipped 
its beak into the mud, lifted its head back, and vigor- 
ously shook its head from side to side several times. 
The jays flew from the pond into the forest canopy 
for several minutes before returning and repeating 
the behavior for another 1 — 2 minutes. This behavior 
of Gray Jays returning to perches between or after 
predation episodes has been noted in other observa- 
tions of predation on amphibians and small mammals 
(Gill 1974; Tordoff 1980; Beiswenger 1981; Strick- 
land and Ouellet 1993; Pilliod 2002). Our inspection 
of the muddy depression after the jays departed the 
vicinity revealed about 24 live Long-toed Salamander 
larvae. Salamanders were similarly sized (ca. 50 mm 
total length) and had full or partial gills. The tiny 
depression they occupied contained saturated mud but 
lacked surface water, macrophytic vegetation or any 
other visible material. We observed Gray Jay-sized 
tracks at other muddy locations within the basin of the 
pond. No other aggregations of stranded salamander 
larvae were found. 

During a 70-min return visit on the afternoon of 12 
October 2003, we observed no Gray Jays or Long-toed 


29] 


202 


Salamanders in or near the same pond perimeter. How- 
ever, we found recent raven and Gray Jay-sized tracks 
at the former saturated depression (which had become 
flooded after an early snow) and along the margins of 
a separate neighboring pond, which also held shallow 
water. We observed no Gray Jays during a 90-min revis- 
it on the afternoon of 21 October 2003, but fresh corvid- 
sized tracks were common in the muddy pond basin. 

The foraging behavior of the Gray Jays, the abun- 
dance of Long-toed Salamanders, and the lack of sur- 
face water and alternative food items in the small 
depression lead us to conclude that the Gray Jays 
were consuming Long-toed Salamanders. Our obser- 
vations are consistent with other descriptions of west- 
ern corvids as opportunistic predators, and suggest 
that some salamander and anuran larvae are suitable 
prey for jays. Unlike some pond-breeding amphibians 
(see Kats et al. 1988), Long-toed Salamander larvae 
appear to be palatable to a variety of predators (Pil- 
liod and Fronzuto 2005). Garter snakes (Thamnophis 
sp.) (Ferguson 1961), non-native fish (Tyler et al. 1998; 
Monello and Wright 2001), Northwestern Salamanders 
(A. gracile) (Hoffman and Larson 1999), conspecific 
larvae (Walls et al. 1993), giant water bugs (Letho- 
cerus sp.) (CAP, personal observation), and dytiscid 
water beetles (Pilliod and Fronzuto 2005) consume 
larval Long-toed Salamanders. 

Gray Jays are common at Crater Lake National Park 
and throughout much of the range of the Long-toed 
Salamander in mountain regions of western North 
America (Farner 1952; Strickland and Ouellet 1993; 
Marshall et al. 2003). Long-toed Salamanders often 
breed in temporary ponds, and larvae are likely to be 
stranded as water levels decline in late summer dur- 
ing dry years (Kezer and Farner 1955). The abun- 
dance of tracks at our observation pond suggests 
Gray Jays or other similarly sized birds visited this 
pond frequently. Long-toed Salamanders and other 
palatable amphibians may represent a seasonally impor- 
tant food resource to opportunistic corvid predators such 
as Gray Jays. 


Acknowledgments 

We thank B. McCreary, D. Olson, D. Pilliod, and 
A. J. Erskine for helpful reviews of the manuscript. 
The National Park Service Inventory and Monitoring 
Program provided support for portions of this research. 


Literature Cited 

Beiswenger, R. E. 1981. Predation by Gray Jays on aggre- 
gating tadpoles of the Boreal Toad (Bufo boreas). Copeia 
1981: 459-460. 

Bent, A. C. 1946. Life histories of North American jays, crows 
and titmice. Smithsonian Institute, United States National 
Museum Bulletin 191. 495 pages. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Brothers, D. R. 1994. Bufo boreas (Western toad). Predation. 
Herpetological Review 25: 117. 

Corn, P. S. 1993. Bufo boreas (Boreal toad). Predation. Her- 
petological Review 24: 57. 

Farner, D. S. 1952. The birds of Crater Lake National Park. 
University of Kansas Press. Lawrence, Kansas. 187 pages. 

Ferguson, D. E. 1961. The geographic variation of Ambystoma 
macrodactylum Baird, with description of two new sub- 
species. American Midland Naturalist 65: 311-338. 

Gill, D. 1974. The Gray Jay as a predator of small mammals. 
Canadian Field Naturalist 88: 370-371. 

Hoffman, R. L., and G. L. Larson. 1999. Ambystoma gracile 
(Northwestern Salamander). Predation and cannibalism. 
Herpetological Review 30: 159. : 

Kats, L. B., J. W. Petranka, and A. Sih. 1988. Antipredator — 
defenses and the persistence of amphibian larvae with fishes. 
Ecology 69: 1865-1870. 

Kezer, J., and D. S. Farner. 1955. Life history patterns of the 
salamander Ambystoma macrodactylum in the high Cascade 
mountains of southern Oregon. Copeia 1955: 127-131. 

Marshall, D. B., M. G. Hunter, and A. L. Contreras. 2003. 
Birds of Oregon: a general reference. Oregon State Uni- 
versity Press, Corvallis, Oregon. 768 pages. 

Monello, R. J., and R. G. Wright. 2001. Predation by Goldfish 
(Carassius auratus) on eggs and larvae of the Eastern 
Long-toed Salamander (Ambystoma macrodactylum colum- 
bianum). Journal of Herpetology 35: 350-353. ) 

Olson, D. H. 1989. Predation on breeding Western Toads — 
(Bufo boreas). Copeia 1989: 391-397. 

Pilliod, D. S. 2002. Clark’s Nutcracker (Nucifraga colum- 
biana) predation on tadpoles of the Columbia Spotted Frog — 
(Rana luteiventris). Northwestern Naturalist 83: 59-61. 

Pilliod, D. S., and J. A. Fronzuto. 2005. Ambystoma 
macrodactylum, Long-toed Salamander. Pages 617-621 
in Amphibian declines: the conservation status of United 
States species. Edited by M. Lannoo, University of Cali- 
fornia Press, Berkeley, California. 

Strickland, D., and H. Ouellet. 1993. Gray Jay. Birds of North | 
America 40. Academy of Natural Sciences and American | 
Ornithological Union. Philadelphia, Pennsylvania. 24 pages. 

Tordoff, W. 1980. Selective predation of Gray Jays, Perisoreus | 
canadensis, upon Boreal Chorus Frogs, Pseudacris trise-_ | 
riata. Evolution 34: 1004-1008. | 

Turner, F. B. 1960. Population structure and dynamics of the — 
Western Spotted Frog, Rana p. pretiosa Baird and Girard, 
in Yellowstone Park, Wyoming. Ecological Monographs | 
30: 251-278. | 

Tyler, T. J., W. J. Liss, R. L. Hoffman, and L. M. Ganio. | 
1998. Experimental analysis of trout effects on survival, | 
growth, and habitat use of two species of Ambystomatid | 
salamanders. Journal of Herpetology 32: 345-349. | 

Walls, S. C., J. J. Beatty, B. N. Tissot, D. G. Hokit, and A. | 
R. Blaustein. 1993. Morphological variation and cannibal- | 
ism in a larval salamander (Ambystoma macrodactylum { 
columbianum). Canadian Journal of Zoology 71: 1543- 
L551. | 


Received 18 February 2004 
Accepted 22 March 2005 


2005 


NOTES 


293 


Territorial Behavior in Belted Kingfishers, Ceryle alcyon, 


During Fall Migration 


MICHAEL J. HAMAS 


Department of Biology, Central Michigan University, Mount. Pleasant, Michigan 48859 USA 


Hamas, Michael J. 2005. Territorial behavior in Belted Kingfishers, Ceryle alcyon, during fall migration. Canadian Field- 


Naturalist 119(2): 293-294. 


The Belted Kingfisher (Ceryle alcyon) exhibits territorial behavior during breeding and on the wintering grounds. However, 
behavior during migration is poorly documented. Here, I report on kingfishers exhibiting territorial behavior during fall 
migration along the shoreline of northern Lake Huron, Michigan. 


Key Words: Belted Kingfisher, Ceryle alcyon, territorial behavior, migration, stopover, Lake Huron, Michigan. 


The Belted Kingfisher (Ceryle alcyon) is widely dis- 
tributed throughout temperate North America where 
breeding populations withdraw from northern latitudes 
and migrate south for the winter. Fall migration 1s ap- 
parent by mid-September, when birds begin to congre- 
gate along major waterways, and continues through 
November (Hamas 1994). Kingfishers are not known 
to travel in aggregated flocks, but in the Great Lakes 
region, Salyer and Lagler (1946) observed a continuous 
progression of individuals moving southward along 
the shoreline of Lake Michigan during October. 

Like other migrants, kingfishers should seek stop- 
over sites providing adequate resources that enable 
birds to continue their migration. Restricted to a diet 
that consists primarily of fishes and aquatic inverte- 
brates, kingfishers’ ability to find food during migra- 
tion depends on clear water, in which prey are easily 
detected. Even wave action on lakes may determine 
where birds occur (Prose 1985). Thus, morphology of 
lacustrine shorelines may influence habitat selection 
by kingfishers at any time of year. 

From 30 August until 23 September 2003, I observed 
solitary kingfishers along approximately 20 km of Lake 
Huron shoreline between Cedarville and DeTour Vil- 
lage in eastern upper Michigan. Forested peninsulas 
and intervening bays characterize the rocky shoreline 
where numerous emergent limestone boulders are errat- 
ically scattered in the shallow nearshore waters. With 
Great Lakes water levels at or near historical lows, 
kingfishers were unable to forage from tree branches 
overhanging water. Instead, birds either hovered over 
open water, or perched on emergent rocks from which 
they would dive for prey. 

Kingfishers occurred at five different bays along the 
shoreline where diurnal variations in wind speed and 
direction sometimes generated waves that forced birds 
to move elsewhere. At Dudley Bay, however, which is 
protected from Lake Huron by a barrier island, wave 
action was minimal. No kingfishers occupied the bay 
during the breeding season, but three male kingfishers, 
two adults and one immature, partitioned the bay into 
contiguous territories which they defended for 11 con- 


secutive days before departing. The birds were not 
marked, but individuals observed fishing from the same 
wooden dock or from the same exposed rocks daily were 
likely the same birds. 

Agonistic encounters between birds included five 
chases observed on different days and an attack. Each 
interaction was accompanied by shrill rattling or 
screams, vocalizations typical of territorial disputes 
(Davis 1988). Observations were made from 06:15 — 
07:30 EST when birds were likely to be foraging. 
Chases were initiated in response to an incursion by a 
conspecific into a portion of the bay occupied by anoth- 
er kingfisher. After driving an intruder from its territory, 
a pursuer would often return to the same fishing perch 
it occupied prior to the chase. Lincoln (1924) noted 
similar behavior in migrant kingfishers responding to 
humans. Sometimes, chases between birds over the tops 
of exposed boulders were saltatory. One bird repeat- 
edly pursued an intruder which retreated to a nearby 
boulder, only to be pursued and displaced again. 

The single attack occurred when an intruder being 
chased by another kingfisher retreated into its territory 
and landed on a dock. With its beak directed downward, 
the pursuer hovered less than | m over the perched bird 
which remained stationary, its crest erected and head 
thrown back. During the encounter, both birds were 
continuously emitting shrill screams. The pursuer then 
landed less than 2 m from the perched bird, and the two 
remained face to face, crests erected and vocalizing for 
nearly three minutes. The pursuer then returned to the 
portion of the bay from which the chase had originated. 

Agonistic intraspecific interactions exhibited by king- 
fishers appear to occur throughout the year suggesting 
that continuous access to adequate prey is requisite and 
promotes territoriality wherever birds occur. During 
breeding, fishing areas are aggressively defended by 
both males and females, and overwintering territories 
are defended by solitary birds of all age classes, but may 
be temporarily abandoned during inclement weather 
(Davis 1980). 

Although stopover territories are ephemeral, monop- 
olizing access to prey is likely to enhance migrants’ 


294 


foraging efficiency. At Dudley Bay none of the birds 
that I observed was marked, but 10 regurgitated pellets 
collected daily at the dock were likely from the same 
bird. Each pellet contained abundant fragments of cray- 
fish exoskeleton in addition to some fish bones suggest- 
ing that kingfishers took prey that was most accessible 
and perhaps most abundant in the shallow waters of the 
bay. Knowing where to find food and excluding com- 
petitors in unfamiliar places are likely to enhance pros- 
pects for a successful migration (Moore 1999). 


Acknowledgments . 

I am grateful to Marilyn Twining for lodging at Dud- 
ley Bay and her ongoing support for avian research 
along the shoreline of northern Lake Huron. 


Literature Cited 

Davis, W. J. 1980. The Belted Kingfisher, Megaceryle alcyon: 
Its ecology and territoriality. M.Sc. thesis, University of 
Cincinnati, Cincinnati, Ohio. 100 pages. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Davis, W. J. 1988. Acoustic communication in the Belted 
Kingfisher: an example of temporal coding. Behaviour 106: 
1-24. 

Hamas, M. J. 1994. Belted Kingfisher (Ceryle alcyon). In 
The Birds of North America, (84). Edited by A. Poole and 
F. Gill. The Academy of Natural Sciences, Philadelphia: 
The American Ornithologists’ Union, Washington, D.C. 

Lincoln, F. C. 1924. A “territory” note on the Belted King- 
fisher. Wilson Bulletin 36: 113-115. 

Moore, F. R. 1999. Neotropical migrants and the Gulf of 
Mexico: the cheniers of Louisiana and stopover ecology. 
In Gatherings of Angels: Migratory Birds and their Ecol- 
ogy. Edited by K. P. Able. Cornell University Press, Itha- 
ca. 193 pages. 

Prose. B. L. 1985. Habitat suitability index models: Belted 
Kingfisher. U.S. Fish and Wildlife Service Biological 
Report (82). 22 pages. 

Salyer, J. C., and K. F. Lagler. 1946. The Eastern Belted 
Kingfisher, Megaceryle alcyon alcyon (Linnaeus) in rela- 
tion to fish management. Transactions of the American 
Fisheries Society 76: 97-117. 


Received 8 August 2004 
Accepted 26 February 2005 


First Record of Age 0+ Atlantic Whitefish, Coregonus huntsmani, 


from the Wild 


DAN J. HASSELMAN!: 3, PHILLIP LONGUE?, and Rop G. BRADFORD? 


'Biology Department, Acadia University, Wolfville, Nova Scotia B4P 2R6 Canada 
"Department of Fisheries and Oceans; Bedford Institute of Oceanography, P. O. Box 1006, 1 Challenger Drive, Dartmouth, 


Nova Scotia B2Y 4A2 Canada 


>Corresponding author (Biology Department, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada 


Hasselman, Dan J., Phillip Longue, and Rod G. Bradford. 2005. First record of age 0+ Atlantic Whitefish, Coregonus huntsmani, 
from the wild. Canadian Field-Naturalist 119(2): 294-295. 


A small school of juvenile Atlantic Whitefish (~20-30 individuals) were observed in the littoral zone of Hebb Lake (Petite 
Riviere watershed), Nova Scotia in June 2000. Of these, a single individual was captured with a beach seine, and confirmed 
to be a young-of-the-year specimen. This is the first documented report of the occurrence of wild Atlantic Whitefish juveniles. 
Occupation of the littoral zone into early summer by young-of-the-year Atlantic Whitefish may have implications for predation 


by invasive Smallmouth Bass. 


Key Words: Atlantic Whitefish, Coregonus huntsmani, wild juvenile, first record, Hebb Lake, Petite Riviere watershed, 


Nova Scotia. 


On 26 June 2000, during daylight experimental angl- 
ing on Hebb Lake (Petite Riviere watershed, Nova 
Scotia; N44°21'00; W64°34'00) to assess susceptibility 
of Atlantic Whitefish (Coregonus huntsmani Scott, 
1987) to incidental capture, a school of small fish were 
observed in pursuit of an ~8cm in length floating im1- 
tation minnow during its retrieval to the shore from a 
cast destination of approximately ~3m depth. Although 
none of these fish were hooked, they remained near 
the shore for a sufficient period of time to determine 
that they were of similar size, silvery in colouration, 
and in possession of an adipose fin. A single Atlantic 
Whitefish (8.4 cm total length; 4.8 g total weight) was 
captured after re-casting the lure, and drawing a small- 
meshed beach seine behind and across the line of 


retrieval to bar escape. The specimen was aged (scale 
method), determined to be a young-of-the-year (y-o-y) 
specimen, and is archived at the Nova Scotia Museum 
of Natural History (Catalogue Number: NSM 85368). 

These are the first observations of juvenile Atlantic 
Whitefish in the wild, as none were available to support 
previous taxonomic descriptions (Smith and Todd 
1992; Edge et al. 1991; Piers 1927; Huntsman 1922) 
or ecological assessments (Edge and Gilhen 2001; Edge 
1984, 1987). Recent (May-August, 2000) systematic 
temporal and spatial sampling with baited minnow pots, 
fyke nets, and beach-seines failed to collect a single 
specimen, although fish species known to frequent the 
shallow (<Im) littoral zone of most Nova Scotia lakes 
(i.e., American Eel (Anguilla rostrata), Banded Killifish 


2005 


(Fundulus diaphanus), Brown Bullhead (Ameiurus neb- 
ulosus), Creek Chub (Semotilus atromaculatus), Golden 
Shiner (Notemigonus crysoleucas), White Sucker (Cat- 
ostomus commersoni), White Perch (Morone ameri- 
cana), and Yellow Perch (Perca flavescens)) were 
captured (R. G. Bradford, unpublished data). Similar 
findings were previously confirmed by Edge (1987). 
Irrespective of the rarity of known occurrences of 
juvenile Atlantic Whitefish within inshore areas of 
shallow—moderate depth, there are indications that the 
species is not precluded from occupation of zones of 
warmer water temperatures. Young-of-the-year Atlantic 
Whitefish reared in captivity routinely tolerate summer 
temperatures of up to 24°C (John Whitelaw, Depart- 
ment of Fisheries and Oceans, Mersey Biodiversity 
Centre, Milton, Nova Scotia, personal communication). 
Recorded June water temperatures at 1 m depth in Hebb 
Lake are a comparable 19.9° + 3.2°C (mean + s.d.), and 
range from 14.2° to 26.1°C (R. G. Bradford, unpub- 
lished data). Young coregonids generally tolerate high- 
er temperatures than do their adult counterparts (Fry 
1937). Therefore, the presence of y-o-y Atlantic White- 
fish in lacustrine habitat shallower than the depth of 
the summer hypolimnion cannot be discounted. 
These observations have significance as an indication 
that small-bodied Atlantic Whitefish may be susceptible 
to direct predation by Smallmouth Bass (Micropterus 
dolomieu), a non-native invasive species illegally intro- 
duced into the upper Petite Riviere drainage during the 
late 1990s (J. Gilhen, Nova Scotia Museum of Natural 
History, personal communication; Atlantic Whitefish 
Conservation and Recovery Team 2004). Smallmouth 
Bass now reproduce successfully in Minamkeak Lake, 
the upper most of three lakes that, when combined, total 
no more than 16 km? of aquatic habitat, and represents 
the global distribution of Atlantic Whitefish (Bradford 
et al. in preparation). Colonization of the other two 
lakes by smallmouth bass is probable in light of the 
experience with this aquatic invasive species else- 
where in North America (Vander Zanden et al. 1999, 
2004). Occurrence of y-o-y Atlantic Whitefish in the 
limnetic zone during the seasons of active foraging by 
Smallmouth Bass may therefore be a matter of impor- 
tance to the survival of the species, but further study is 
required to determine the validity of this supposition. 


Acknowledgments 

Observations occurred during field activities in sup- 
port of the Atlantic Whitefish recovery strategy. Fund- 
ing was provided by the Canadian Department of Fish- 
eries and Oceans. 


NOTES 


295 


Documents Cited 

Bradford, R. G., D. Longard, and P. Longue. In prepration 
Status, trend, and recovery considerations in support of an 
allowable harm assessment for Atlantic whitefish (Core- 
gonus huntsmani), Canadian Stock Assessment Secre- 
tariat Research Document 2004, Department of Fisheries 
and Oceans, Dartmouth, Nova Scotia. 


Literature Cited 

Atlantic Whitefish Conservation and Recovery Team. 
2004. National recovery strategy for the Atlantic Whitefish 
(Coregonus huntsmani). Department of Fisheries and 
Oceans. 2004. Halifax, pages 34. 

Fry, F. E. J. 1937. The summer migration of the cisco, Leu- 
cichthys artedii (Le Sueur), in Lake Nipissing, Ontario. 
Publications of the Ontario Fisheries Research Laboratory 
55: 9-91. 

Edge, T. A. 1984. Preliminary status of the Acadian whitefish, 
Coregonus canadensis, 1n southern Nova Scotia. Canadian 
Field-Naturalist 98: 86-90. 

Edge, T. A. 1987. The systematics, distribution, ecology and 
zoogeography of the endangered Acadian whitefish, Core- 
gonus canadensis Scott, 1967, in Nova Scotia, Canada. 
M.Sc. University of Ottawa. 

Edge, T. A., D. E. McAllister, and S. U. Qadri. 1991. 
Meristic and morphometric variation between the endan- 
gered Acadian whitefish, Coregonus huntsmani, and the 
lake whitefish, Coregonus clupeaformis, in the Canadian 
Maritime Provinces and the state of Maine, USA. Canadian 
Journal of Fisheries and Aquatic Sciences 48: 2140-2151. 

Edge, T. A., J. Gilhen. 2001. Updated status report on the en- 
dangered Atlantic Whitefish, Coregonus huntsmani. Cana- 
dian Field-Naturalist 115: 635-651. 

Huntsman, A. J. 1922. The fishes of the Bay of Fundy. 
Contribution to Canadian Biology 3: 49-72. 

Piers, H. 1927. Coregonus labridoricus, the Sault Whitefish, 
an interesting addition to the freshwater fish fauna of Nova 
Scotia. Transactions of the Nova Scotia Institute of Science 
16: 92-95. 

Smith, G. R., and T. N. Todd. 1992. Morphological cladistic 
study of coregonine fishes. Polish Archives of Hydrobi- 
ology 39: 479-490. 

Vander Zanden, M. J., J. M., Casselman, and J. B. Ras- 
mussen. 1999. Stable isotope evidence for the food web 
consequences of species invasions in lakes. Nature 401: 
464-467. 

Vander Zanden, M. J., J. D. Olden, J. H. Thorne, and N. 
E. Mandrak. 2004. Predicting occurrences and impacts 
of smallmouth bass introductions in north temperate lakes. 
Ecological Applications 14: 132-148. 


Received 2 November 2004 
Accepted 12 March 2005 


296 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Passage Through a Small Drainage Culvert by Mule Deer, 
Odocoilus hemionus, and Other Mammals 


ALISSA KRAWCHUK!, KARL W. LARSEN!, RICHARD D. WEIR2, AND HELEN Davis? 


‘Department of Natural Resource Sciences, Thompson Rivers University, PO Box 3010, Kamloops, British Columbia V2C 5N3 


Canada 


Artemis Wildlife Consultants, 4515 Hullcar Road, Armstrong, British Columbia VOE 1B4 Canada 


Krawchuk, Alissa, Karl W. Larsen, Richard D. Weir, and Helen Davis. 2005. Passage through a small drainage culvert by 
Mule Deer, Odocoilus hemionus, and other mammals. Canadian Field-Naturalist 119(2): 296-298. 


Cameras with infra-red triggers were used to monitor the passage of wildlife through underground passages that ran under a 
major highway and railway. Several species of mammals were detected traveling through the passages; of particular interest 
was the movement of Mule Deer (Odocoileus hemionus) through a relatively small culvert that would not have been predicted 


to see usage by these animals. 


Key Words: road crossing, Mule Deer, Odocoilus hemionus, Black Bear, Ursus americanus, road ecology, underpass, highway 


crossing. 


Numerous types of crossing structures exist to aid in 
the movement of wildlife across highways and rail- 
ways. Some of these structures, such as drainage cul- 
verts, cattle underpasses and human underpasses, are 
not specifically designed for use by wildlife, yet they 
are ubiquitous features associated with highways, and 
are far more common than underpasses and overpasses 
specifically designed for wildlife. To this end, their 
potential and realized roles in reducing negative road 
effects on wildlife need to be explored. Culverts, for 
example, are known to be used by numerous types of 
wildlife including small mammals, reptiles, amphib- 
ians and large carnivores (Yanes et al. 1995; Rod- 
riguez et al. 1996; Clevenger et al. 2001). 

Interest in the ability of crossing structures to lower 
deer mortality on highways grew in the 1970s in res- 
ponse to large numbers of these animals being struck 
by vehicles in the United States (Puglisi et al. 1974; 
Allen and Cullough 1976). In particular, Reed et al. 
(1975, 1979) studied the use of underpasses by Mule 
Deer, Odocoilus hemionus, and as a result, the suitabili- 
ty of crossing structures for deer was well document- 
ed. Reed et al. (1975, 1979) identified the openness ratio 
(width*height/length) of these structures as an impor- 
tant factor influencing deer use. Their work suggested 
that structures with openness ratios of less than 0.6 
would not be utilized by deer. This minimum guideline 
is somewhat difficult to test, as structures monitored for 
wildlife passage often are much larger (Yanes et al. 
1995; Foster and Humphry 1995). One recent study 
that examined small crossings found that deer would use 
culverts as small as 4.2 m wide, 3.5 m tall, and 96.1 m 
long (openness ratio of 0.15, Clevenger and Waltho 
2000). In the same study, deer also were found to use 
underpasses as large as 14.9 m wide, 3.2 m tall and 
38.0 m long (openness ratio of 1.25). 

We conducted a small, localized study to document 
whether or not wildlife was using various passages 
under major transportation corridors in the region sur- 
rounding Kamloops, British Columbia. Herein we re- 


port on our results, particularly the repeated use of an 
unusually small culvert by deer. 


Methods 

This study took place in the fall of 2003. We chose 
three crossing structures (two drainage culverts and one 
cattle underpass) east of Kamloops, British Columbia 
(119°52'W, 50°39'N) to monitor with infra-red sensor 
cameras. Each of these three crossings extends under 
both the Trans-Canada Highway and the Canadian 
Pacific Railway. The former is the major transportation 
corridor for motorists traveling through the Kamloops 
region. Monthly average daily traffic on this road for 
September to November ranges between 10 000 and 
15 000 vehicles per day (British Columbia Ministry of 
Transportation, Transportation Information Manage- 
ment System data [1997-1999]). At two of the three 
sites we monitored (Site 1 and 2), the highway con- 
sisted of four lanes for at least 1 km in either direc- 
tion. The dimensions of the crossing structures at the 
three sites are summarized in Table | (see Figure 1). 

We used camera units (MacHutchon et al. 1998) 
constructed from 35 mm cameras with autofocus, wide 
angle lens, autoflash and date/time function (Olympus 
Af-1 Twin, Japan). These units were linked to passive 
infra-red sensors (Sureshot 6257-SPDT, Sentrol, USA) 
and hooked through a converter and an on/off switch 
to a 12 volt battery. The sensors were sensitive enough 
to be triggered by very small movements such as a 
breeze gently moving plants or a small animal passing 
by, if the movement was close enough to the camera 
(within 20 m). The entire assembly was housed in a 
modified ammunition box that had been painted black. 

The camera boxes were attached to right-angle 
brackets that allowed mounting on trees or fenceposts. 
The boxes also could be anchored directly to the ground, 
in situations where suitable mounting support was un- 
available. Due to site characteristics and a limited 
availability of mounting structures, it was not always 
possible to position the cameras directly facing the 


2005 


NOTES 


207 


TABLE |. Crossing occurrence of wildlife in culverts, as detected by infra-red sensors and cameras near Kamloops, British 


Columbia, autumn 2003. 


Passage Passage Passage Openess Ratio Species detected Number of 
Location Width Height Length (width*height detected 
(m) (m) (m) /length) crossings 
Site 1* ea LS 30 0.11 Black Bear 20 
Mule Deer 6 
Site 2" 10 5.0 40 0.88 Black Bear 4 
Raccoon 2 
Site 3*** ee i 30 0.05 none detected - 


* camera operational August 21 to November 13, 2003 (Figure 1) 


** camera operational August 30 to November 13, 2003 
*** camera operational September 5 to November 13, 2003 


culverts. We also tried to position the cameras so that 
the flash would not go off directly in the eyes of any 
wildlife using the structure, thereby startling them and 
possibly discouraging use. 

We visited each site and its camera station every few 
days, depending on the frequency of pictures taken and 
the battery strength. A spare battery allowed for rota- 
tion between the three cameras with minimal camera 
downtime. At each camera check, the battery strength 
and the number of exposures were noted. The presence 
of animal tracks or sign around the camera station also 
was noted, to confirm that the cameras were not miss- 


ing animals. From the developed photos, we counted 
the number of animals of each species detected at each 
camera. When the same animal was seen in a sequence 
of photos, it was only counted as one crossing event. 


Results 

Despite the brevity of the study and the small num- 
ber of locations monitored, we were successful at de- 
tecting use of the crossing structures at two of three 
camera sites. Table 1 summarizes the native species 
documented using the crossing structures. Non-wildlife 
use detected by the cameras included humans, cattle 


‘< 


ib ES OE 


Bat 


FiGuRE |. Drainage culvert at Site 1, used repeatedly by Mule Deer to travel under the Trans-Canada Highway near Kamloops, 
British Columbia (dimensions of culvert: 2.1 m wide, 1.5 m tall, 30 m long). 


298 


and housecats. No photo of wildlife crossing was ob- 
tained at Site 3; however, a faulty flash mechanism on 
the camera at this site rendered all night-time photos 
indecipherable, so we simply may have failed to detect 
animals using this culvert for passage. 

Mule Deer were detected traveling through the Site 
1 culvert that had an openness ratio of only 0.11 (Ta- 
ble 1). The camera at this site recorded Mule Deer on 
six different occasions over the 10-week study. This 
culvert had a dirt substrate, was surrounded on either 
side by vegetation, and was relatively far from human 
activity. Deer were not detected at Site 2, however, a 
road-killed deer was found within 50 m of the culvert 
during the study period, suggesting deer were in the 
vicinity. Deer also were not detected at Site 3, possibly 
due in part to the faulty flash (see above), but deer 
tracks were seen in the area of the underpass during 
the study period. 


Discussion 

Our general observations of Black Bear, Ursus 
Americanus, Raccoon, Procyon lotor, and Mule Deer 
passing through the monitored culverts are in keeping 
with results from other studies (e.g., Clevenger and 
Waltho, 2000; LaPoint et al. 2003). However, of par- 
ticular interest is the use of the drainage culvert at Site 
| by Mule Deer, as previous work has suggested that 
ungulates are reluctant to use structures less than 7 m 
wide or 2.4 m high, or with an openness ratio of less 
than 0.6 (Reed et al. 1975; Reed et al. 1979; Yanes et 
al. 1995). This is substantially greater than the ratio for 
the culvert at Site | in this study. However, Clevenger 
and Waltho (2000) recently reported on the use by deer 
of an underpass with an openness ratio of 0.15 in Banff, 
which together with our study supports the notion that 
deer may be more plastic in their use of crossing struc- 
tures than previously thought. 

Our failure to detect deer crossing at Sites 2 and 3 
(even though evidence showed deer were in the area) 
likely is due in part to our small sampling period, and 
a host of other potential factors (Rodriguez et al. 1996, 
Clevenger et al. 2001). Unlike Site 1, the culverts at 
Site 2 and 3 lacked a large amount of surrounding 
natural vegetation, a feature found to increase the use 
of crossings by wildlife (Yanes et al. 1995). The natural 
dirt substrate on the bottom of the culvert at Sitel also 
may have encouraged its use by deer, as a substrate of 
soil and detritus has been reported to be less hostile 
to wildlife (Reed et al. 1975; Yanes et al. 1995). Fur- 
ther, it has been suggested that elevated noise levels 
associated with human activity deter wildlife from 
utilizing culverts close to such areas (Clevenger et al. 
2000), and the Site | culvert, where deer were detect- 
ed, was relatively far away from human activity. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Despite the limitations of our study, we document- 
ed the passage of Mule Deer under a major highway, 
through a culvert that would not have been predicted 
to show usage by these animals. Our observations sug- 
gest further research is required on the use or avoid- 
ance of culverts and other structures not specifically 
designed for wildlife. 


Acknowledgments 

The Smith family graciously allowed us to set up a 
monitoring camera on their property near Pritchard, 
British Columbia. Thanks to E. Lofroth and T. Hamil- 
ton for facilitating this project. This study was fund- 
ed by the Habitat Conservation Trust Fund of British 
Columbia. 


Literature Cited 

Allen, R. E., and D. R. McCullough. 1976. Deer-car acci- 
dents in southern Michigan. Journal of Wildlife Manage- 
ment 40: 317-325. 

Clevenger, A. P., and N. Waltho. 2000. Factors influencing 
the effectiveness of wildlife underpasses in Banff National 
Park, Alberta, Canada. Conservation Biology 14: 47-56. 

Clevenger, A. P., B. Chruszcz, and K. Gunson. 2001. 
Drainage culverts as habitat linkages and factors affecting 
passage by mammals. Journal of Applied Ecology 38: 
1340-1349. 

Foster, M. L., and S. R. Humphry. 1995. Use of highway 
underpasses by Florida panthers and other wildlife. Wildlife 
Society Bulletin 23: 95-100. 

LaPoint, S. D., R. W. Kays, and J. C. Ray. 2003. Animals 
crossing the Northway: are existing culverts useful? Adir- 
ondack Journal of Environmental Studies 10: 11-17. 

MacHutchon, A. G., S. Himmer, H. Davis, and M. Gal- 
lagher. 1998. Temporal and spatial activity patterns among 
coastal bear populations. Ursus 10: 539-546. 

Puglisi, M. J., J. S. Lindzey, and E. D. Bellis. 1974. Factors 
associated with highway mortality of white-tail deer. Jour- 
nal of Wildlife Management. 38: 799-807. 

Reed, D. F., T. N. Woodward, and T. M. Pojar. 1975. Be- 
havioural response of Mule deer to a highway underpass. 
Journal of Wildlife Management 39: 361-367. 

Reed, D. F., T. N. Woodward, and T. D. I. Beck. 1979. 
Regional deer-vehicle accident research. U.S. Department 
of Transport Federal Highway Adminstration Report 
FHWA-RD-79-11. National Technical Information Service, 
Springfield, Virginia. 61 pages. 

Rodriguez, A., G. Crema, and M. Delibes. 1996. Use of 
non-wildlife passages across a high speed railway by ter- 
restrial vertebrates. Journal of Applied Ecology 33: 1527- 
1540. 

Yanes, M., J. M.Velasco, and F. Suarez. 1995. Permeability 
of roads and railways to vertebrates: the importance of 
culverts. Biological Conservation 71: 217-222. 


Received 27 September 2004 
Accepted 6 February 2005 


Book Reviews 


ZOOLOGY 


Birds of Azerbaijan 
By Michael Patrikeev. 2004. Pensoft Publishers, Geo Milev 


Street 13a, 1111 Sofia, Bulgaria. 500 pages, U.S. $177.50 
Cloth. 


Several years ago I took a small map and shaded 
the countries for which you could purchase a good 
bird guide. There were two key gaps. One zone ran 
from eastern Turkey to Afghanistan (and the other 
was Brazil). The best books you could get for this 
former region (the Caucasus) was the Birds of Russia 
by Flint Boehme, Kostin and Kuznetsor or Birds of 
the Soviet Union by Dementiev and Gladkov (both 
nearly 50 years old). This new book covers the 372 
species that have been recorded in the Republic of 
Azerbaijan and fills in some of that area with poor 
coverage. 

Azerbaijan lies on the western shore of the Caspian 
Sea, to the north of Iran and east of Turkey. A southern 
section of Armenia clips off a small enclave called 
Noxcivan. It has a varied topography with the conse- 
quent diversity of bird species. This book covers this 
troubled region. There were wars in the 1800s with 
imperial Russia, civil war between 1917-1920, attacks 
by the Turks from the 1950s onwards, and still the area 
has an ongoing dispute with neighbouring Armenia. 
The struggles, along with changing economic and 
political status has led to some discontinuity in the 
study of birds. Separation from the USSR in 1991 lost 
the services of the Russian science community and this 
lack of continuous research shows in the data cited. 
Much of the information used by the author is from 
before the mid-1990s. There are some later refer- 
ences, but these are much fewer. Separation from the 
former USSR also caused economic difficulties from 
which the country has yet to emerge. 

Birds of Azerbaijan is a distributional atlas, not a 
field guide. The author has compiled data from a large 
variety of sources, much from the “Russian” era. For 
example the data on Mallard is primarily prior to 1996. 
[We have seen with our work on the Breeding Bird 
Atlas of Ontario what an amazing difference a gap of 
ten years in the data can make]. The author has added 
his own extensive observations taken between 1970 


Birds of New Brunswick: An Annotated List 


By David S. Christie, Brian E. Dalzell, Marcel David, Robert 
Doiron, Donald G. Gibson, Mike H. Lushington, Peter 
A. Pearce, Stuart I. Tingley, and James G. Wilson. 2004. 
New Brunswick Museum Monographic Series (Natural 
Science) Number 10, 277 Douglas Avenue, Saint John, 


and 199]. The story is quite gloomy. For example, 
many thousands of Red-breasted Geese were seen in 
the 1950s, but the species is considered virtually 
extinct now. Even the Eurasian Coot has dropped 
from millions to tens of thousands. The reasons for 
these changes include severe air, soil, and water pol- 
lution. Soil pollution results from oil spills, DDT and 
defoliants used in the production of cotton, making 
the Caspian Sea one the most ecologically degraded 
area in the world. Less gloomy, but equally distress- 
ing is the lack of current information on some species, 
for example Red Kite. 

It is against this background that the author has 
done a sterling job of pulling the known information 
on this county’s birds into a logical, readable text. 
Each species account has status, distribution, popula- 
tion size, migratory movements, breeding informa- 
tion, diet and mortality. The English names are used 
throughout, while Azaeri names are added for many 
species. A respectably-sized distribution map has codes 
for summer, winter, nesting, etc. for most species. 
The author gives an annotated bird list and describes 
important bird areas, places for colonial birds, water- 
fowl wintering areas, and the influence of cold winters 
and oil pollution. There are 78 photographs, of which 
one third show habitat, one third are bird photos and 
the rest are of eggs and nests. The habitat photographs 
show the nine basic types of landscapes from alpine 
meadows in mountains to semi-desert and wetland. But 
he does not illustrate the rust and mauve mountains 
of the Caucasus located in the troubled Naxg¢ivan 
Autonomous District nor of any arid badlands. There 
is not full coverage of the 50 Important Bird Areas 
identified by the author. 

Given the way the Azaeri economy has languished 
as regional trade has suffered and the underdevel- 
oped oil production has yet to fulfill its promise, this 
book may be a very important milestone in a contin- 
uing tragic ornithological history. 

Roy JOHN 
2193 Emard Crescent, Ottawa, Ontario KIJ 6K5 Canada 


New Brunswick, E2K 1E5. 2004. 83 pages. [Also avail- 
able in French. | 


This publication appeared in the same series as the 
two previous summaries (hereafter “‘lists”) of bird status 


299 


300 


in New Brunswick (Squires 1952, 1976), and seems 
to serve essentially the same purpose, to update the 
provincial bird list and the status of each species. 
This review considers the new publication’s success 
in achieving its purpose, whether viewed only as an 
update of the earlier lists, or as state-of-the-art repre- 
sentation of each bird species’ status in New Brunswick. 

The subtitle “An annotated list” implies that this 
publication was planned as a less ambitious publica- 
tion than its predecessors, but that may be a quibble. 
The earlier books were also annotated lists, though 
including more descriptive “front material” and fairly 
extensive bibliographies. The new species accounts 
generally are shorter than those in the earlier lists, 
omitting most place-names and dates — except for 
species known from very few records. Presumably that 
reflected more complete perspective on status than was 
available earlier, as might be expected; given 30 years’ 
additional data, and their consideration by a panel of 
authors drawn from most parts of the province, it 
may have been redundant to document in detail the 
distribution of each species, many of which occur 
regularly in all or most of the 15 counties. 

Considered only as an updated provincial list, with 
status assessed from existing, largely unstandardized 
data, I judge that this publication will serve most of its 
immediate objectives adequately. Birders will be able 
to find out whether a scarce species has been reported 
previously in New Brunswick and, if so, where and how 
many times. They also can obtain an idea, in very gen- 
eral terms, of how common a regular species is, across 
the province and in the seasons when it is to be ex- 
pected. 

Scientists seeking potential study areas may be 
dissatisfied with the scarcity of specific locations and 
of distributional limits within the province. Few “hot- 
spots” are identified except for very scarce species. 
Admittedly, scientists “from away” probably also had 
difficulty extracting such information from the earlier 
lists, owing to far greater gaps in knowledge then. 

The introduction makes it clear that the present list, 
longer by 61 species than its 1976 predecessor, grew 
by addition largely of vagrants (= “lost birds”). Some 
new species had expanded their ranges, but very few 
of those — Mourning Dove the most obvious example 
— breed widely in New Brunswick as yet. Only two 
species were stated to have declined drastically, though 
other declines were noted in species accounts; my 
recent experience (quite limited geographically) sug- 
gested that some other declines may be as significant 
as those highlighted in the Introduction. 

Perhaps a more important question emerging from 
review of this publication is whether its format was 
suitable for presenting status of all species represen- 
tatively, even if we acknowledge that complete infor- 
mation will never be available? Status involves frequen- 
cy of encounter and density, as well as distribution. 
Most observational effort by birders is notoriously 
uneven, focusing on “hot-spots” and easily accessi- 


THE CANADIAN FIELD-NATURALIST 


Vol. 129 


ble areas — and on misplaced species. Is it reasonable 
to accept that data from such efforts provide the best 
picture available on bird status? 

The status of many breeding birds in New Bruns- 
wick was presented, probably more representatively 
(up to that time), in the maps and population esti- 
mates of the Maritimes Atlas (Erskine 1992). From 
the start of that (breeding) Atlas project, it was recog- 
nized that the major increases in perspective would 
emerge for species that were neither everywhere nor 
with very restricted breeding ranges — and that was 
found to be so. Although “abundance indices” were 
assessed for most species, and extrapolated to total 
breeding populations for many of them, no attempt 
has been made — as yet — to map relative densities of 
breeding species within New Brunswick from Atlas 
data. That would be essential for representative assess- 
ment of status, and it may soon become possible using 
“point-count” data to be collected in the second-gen- 
eration bird atlas in the Maritimes (now in planning) 
[that in Ontario is nearing completion]. Breeding 
species make up most of the provincial list of regu- 
larly occurring birds, and Atlas data should be pre- 
ferred over even up-to-date annotated lists by most 
scientists and many birders. 

The Atlas mapping approach reduced unsampled 
areas greatly — though not completely. It could be used 
effectively only during breeding and wintering sea- 
sons, when birds are fairly sedentary. As yet, no bird 
atlas coverage is available in the Maritimes for winter 
or migration seasons — except for seabirds (which may 
not provide a model useful for other species). Many 
(uncompiled) data exist for other regular species in 
other seasons; although most were collected unsystem- 
atically and with no attempt at comprehensive cover- 
age, it would be possible to produce maps for occur- 
rence and frequency of wintering and migrating birds 
— in this and nearby provinces — that would improve 
greatly on the rather subjective status assessments 
available in check-list form. Until that is done, anno- 
tated lists may be the only readily available summaries 
of bird status in non-breeding seasons, including most 
occurrences of “displaced birds”. 

The new list of New Brunswick birds is a neat, 
compact publication, with a sturdy “ring” binding 
and attractive cover pictures. The front cover features 
an Evening Grosbeak, a colourful and conspicuous 
species that often dominated the New Brunswick bird 
scene in the 1970s — but that, in the past decade, has 
almost disappeared from southeast New Brunswick. 


References: 

Erskine, A. J. 1992. Atlas of breeding birds of the Maritime Provinces. 
Nova Scotia Museum and Nimbus Publishing. x + 270 pages. 

Squires, W. A. 1952. The birds of New Brunswick. Monographic Series 
(4). New Brunswick Museum, Saint John. 164 pages. 

Squires, W. A. 1976. The birds of New Brunswick (Second edition). Mono- 
graphic Series (7). New Brunswick Museum, Saint John. 221 pages. 


A. J. (TONY) ERSKINE 


16 Richardson Street, Sackville, New Brunswick E4L 4H6 
Canada 


2005 


BOOK REVIEWS 


301 


Fishes of the Great Lakes Region Revised Edition 


By Carl L. Hubbs and Karl F. Lagler. Revised by Gerald R. 
Smith. 2004. The University of Michigan Press, Ann Arbor, 
Michigan USA. xvii + 276 pages, plates 1-32. $24.95. 


This book first appeared in 1941 in a different form 
and has had several subsequent editions. It is renowned 
for standardizing the methods for counting scales and 
fin rays and for measuring a fish specimen. This lat- 
est edition adds thirteen newly introduced species, 
plus hybrids between White Bass and Striped Bass, 
and geographical distribution maps for each species. 
It is much more user friendly than the 1964 edition. 
As was intended originally, this book is aimed at stu- 
dents and the ease with which this book can be used 
makes this point clear. 

The Great Lakes region covers 287 770 square miles, 
while the Great Lakes themselves occupy 94 700 square 
miles of this area. Michigan is wholly encompassed by 
the region, as are small portions of other states bor- 
dering the lakes, but Ontario is the largest land area 
within the Great Lakes region so the book has a Cana- 
dian relevance. Within this region there are 28 families, 
70 genera, 161 species and 215 forms of native fish. 
The term “form” is undefined, and seems to be used 
interchangeably with the term “kind”. Presumably sub- 
species is meant. There are 210 species in total. Thir- 
teen species and hybrids have been recently introduced 
by the construction of canals, by stocking, and through 
the dumping of ballast water from international freight- 
ers. Warning is given about the more destructive in- 
vaders’ effects on the fish fauna and the loss of species 
that will result when the full effect is seen. The intro- 
ductory sections of this book go into significant detail 
about the waters of the Great Lakes region, zoogeog- 
raphy, and the effects of postglacial redispersal. 

The information in this book is generally well laid 
out. There is a section on the collection of fishes, on 
the preservation of fishes, and on how to identify these 
preserved fishes. All this information is very conven- 
ient for an amateur naturalist, including the description 
of the extent to which field notes should be made. 
These sections are brief, however, and are intended 
as basic information only. 

There is a key to families and keys to identify 
species within families. The key to families is very 
easy to use, outlining the basic and most prominent 
features, allowing for quick and easy use, even by 
beginners. Not all the keys are together but the page 
references are clear. The species keys are also fairly 
easy to use, and the added line drawings can be helpful 
when identifying vague features but additional key 
features are in a second section after each key. There 
are 62 cyprinid species (carps and minnows) which 
makes it easy to go astray. Perhaps an initial key to 
genera would reduce the chance of error. Some exotics 
_ (Goldfish, Carp) are included in the key but others 
are not (Bighead, Black, Grass and Silver carp). A 
section on these “alien species” appears at the end of 


the key to Cyprinidae and might better be placed at 
the beginning, as would a section on hybridization in 
Cyprinidae. 

Before each key there is a description of the family 
which is limited to physical features, geographical dis- 
tribution, a brief biological outline, and occasionally 
the importance of the family to the fisheries. Species 
introduction dates and effects are included where appli- 
cable. After each key there is a list of all the species 
supplemented by a brief description, including physi- 
cal features, geographical distribution, and what the 
species feeds on. The arrangement of species is not 
alphabetical by common name or by scientific name 
and does not follow the appearance of the species in 
the key, necessitating some searching in larger fami- 
lies. In addition to the descriptions there is a black- 
and-white photograph of the whole fish on the facing 
page as well as small line drawing of heads, spines, 
gill arches, mouths or other key features. The photos 
here, however, are not of the highest quality. In some 
cases, almost all detail is missing. The line drawings 
are small and often have no indication of the key struc- 
tures so necessitating constant flipping between the 
key and the description section. The description has a 
tiny (22 x 16 mm) geographical distribution map. The 
maps are provided as a quick reference only since 
exact locations can scarcely be found. Larger maps are 
provided on the inside of each cover, in significantly 
more detail, allowing for some cross-referencing. On- 
tario and Quebec records of the Chestnut Lamprey 
are not apparent, the Carp and Freshwater Drum are 
not recorded from the area around Ottawa where they 
are common, and the White Bass is mapped for Ottawa 
where it does not occur. Evidently, the maps must be 
taken as a general guide to distribution and cannot 
always be used locally, where most students and nat- 
uralists work. 

At the end of the book there is an appendix of line 
drawings of the larval stages of some of the fish of 
the Great Lakes region. This book is geared towards 
the identification of adult fish and any inquiries as to 
the identification of larvae are directed to consult other 
works. The index includes species names, common 
names and some key words. No glossary is provided 
though its presence would be ideal. While a section 
on anatomy is given at the beginning, it is not useful 
as a quick reference, nor is it complete (for example, 
adipose and falcate are not explained in the book, nor 
are markings well described). The book ends with a 
series of 32 colour plates with several species per plate, 
which are generally useful to flip through to locate a 
species that you are already somewhat familiar with, 
although many of the colours are inaccurate making 
identification by a beginner difficult. 

The page numbering system is a little confusing, 
sometimes at the bottom of the page, sometimes at 
the top, sometimes missing and easily confused with 


302 


the numbering system for species. There are also some 
errors of omission and commission. It is noted that 
scales are not used for ageing mudminnows but does 
not explain why. The number of lamprey species is 
given as about two dozen (the same as in the 1964 
edition) but this number has increased to about 34 
since then. Some scientific names are given as a tri- 
nomial; e.g., Semotilus atromaculatus atromaculatus, 
but no other subspecies are mentioned which makes 
this extensive name unnecessary. 

The list of families provided includes native fish 
only, so to get a comprehensive count of all the dif- 
ferent genera and species in the Great Lakes region 
you would have to look through the entire book. 
Also, upon comparing the present list to the list pro- 
vided in the 1964 edition, the latter includes all fish, 
not just native species. It is therefore not possible, 
without great effort, to find out how many new native 
fish have been discovered in the past forty years. There 
have also been many name changes that have occurred 
since the last edition was published. Entosphenus 
lamotteni has now become Lampetra appendix, Lep- 
isosteus productus has become Lepisosteus oculatus, 
and Pomolobus pseudoharengus has become Alosa 
pseudoharengus, to name a few. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Some of the criticisms of this book mentioned 
here could be addressed by an on-line version. This 
book must be small and therefore concise for use in 
the field and laboratory. An on-line version could 
have larger maps (updated as new information or cor- 
rections are noted), colour photographs of live fish 
which would be of use to field workers, keys seg- 
mented by lake or country (most work is done local- 
ly, a worker in eastern Ontario would not find the 
same suite of species as a worker in Illinois), larger 
illustrations of key characters, key characters embed- 
ded in the key for easier reference, new species could 
be added as discovered, and keys to larval fish devel- 
oped. 

This book is a great tool for use in the lab or field 
by beginners and professionals alike. With some use 
it will lay open flat, preventing the loss of place 
while your hands are busy working on the fish. The 
positive reputation of this book as the book for Great 
Lakes region fishes is long standing and will only 
increase with time. 


KRYSTAL LAPIERRE and BRIAN W. COAD 


Canadian Museum of Nature, Ottawa, Ontario KIP 6P4 
Canada 


Seabirds and Atlantic Canada’s Ship-Source Oil Pollution 


By F. Wiese. 2002. World Wildlife Fund, 245 Eglinton Avenue 
East, Suite 410, Toronto, Ontario M4P 3J1 Canada. 82 pages. 


The public seems to have a love-hate relationship 
with oil. Oil allows for high salaries and contributes 
to civilisation, but it also can create severe pollution. As 
this report shows well, chronic offshore oil ranks among 
the most severe pollution problems in the world. 

“Many people consider Canada to be one of the 
leading nations in environmental conservation in the 
world”. However, the still conservative estimate of 
300 000 dead Canadian seabirds due to chronic oil 
pollution and presented in this report is shocking; to 
say the least. Besides a seabird population issue this 
also a major animal care issue: over 300 000 animal 
individuals are suffering and are dying a gruesome 
death. As this informative report emphasizes, for each 
oiled seabird found in Newfoundland one can assume 
that at least 10 more have died. 

Together with several individuals devoted to the 
issue of marine and oil pollution, author Dr. F. Wiese 
studied seabirds and their oil-related mortality for 
many years. His report on chronic offshore oil pollu- 
tion is structured in two parts: The Problem (11 chap- 
ters) and The Solution (8 chapters); four appendices, a 
list of abbreviations and some references are also given. 
Half of the report deals with OSIRs (Oil Spill Intelli- 
gence Reports 1997-2000), presented in Appendix 4. 
OSIRs are only accessible for few signed-up mem- 
bers, and it is great that this report provides the wider 
public with an opportunity to access this information. 


Besides reporting baseline numbers of seabird mor- 
talities and oil pollution incidents, other highlights of 
this document are presented to a wide audience deal- 
ing with ocean modelling, detectabilities of oiled birds 
on a beach, drift block experiments and emphasizing 
how important such methods are to address the chronic 
offshore oil pollution efficiently and in accurate terms. 

“Most of those in the marine industry carry out their 
operations in a safe and environmentally responsible 
manner.” This statement is somewhat in contrast to the 
fact that oiled birds keep washing up on shorelines 
worldwide which suggests that national legislation and 
international conventions and guidelines are not being 
followed or that they are inefficient. “The illegal dis- 
charge of oil from ships into the world’s oceans is a 
global problem that affects the entire marine ecosys- 
tem”. This calls for a global oiled bird survey; e.g., 
citizen- and volunteer-based marine and beach surveys. 

From this nice report it becomes quickly obvious 
that the history and track-record of chronic oil pollu- 
tion, a by-product of the current civilisation, is not an 
environmental success story. Instead, the current 
progress for trying to keep the (marine) environment 
clean presents more of an international embarrass- 
ment. It is still difficult to understand why the “bur- 
den of proof” for chronic oil pollution is not on the 
industry side. After reading this report and its seabird 
facts one cannot deny that oil pollution equals envi- 
ronmental massmurder. 

It is correct that the Canadian legislation extends 
the enforcement of shipping, environmental, and 


2005 


wildlife law to the 200-mile exclusive economic 
zone. However, some federal legal decisions have 
restricted these laws to the 12-nautical-mile territori- 
al zone. Offshore pollution monitoring flights are 
made by Transport Canada and by the Canadian 
Coastguard; the Department of National Defence and 
Department of Fisheries and Oceans are encouraged 
as well. Although the Canadian Shipping Act, the 
Migratory Bird Convention Act, the Canadian Envi- 
ronmental Protection Act and the Fisheries Act deal 
with oil pollution issues, suspected ships have rarely 
been turned back to a Canadian port for further 
investigations. Only the Migratory Bird Convention 
Act protects migratory seabirds from oil-related 
offences; but so far, only five vessels were charged. 
Knowing that approximately 2500 offshore oil spills 
are reported per year in Atlantic Canada, one gets 
quickly an idea of the issue. Obviously, pollution 
pays ... and as the report convincingly shows, the 
pollutor gets almost awarded due to the competitive 
business advantage when not punished. Atlantic 
Canada is simply the cheapest place to dump bilge 
oil on the Great Circle route between North Atlantic 
and Europe. No doubt, the enforcement needs to be 
stronger in Canada, higher fines are required, and on 
board disposal facilities, increased monitoring, 
increased awareness and other measures are neces- 
sary. Even the European Union uses RADARDSAT 
(SAR), a satellite image product from Canada, in 
order to trace and to monitor offshore oi! pollution. 

As Wiese’s WWE report presents, Canada does not 
really have a national standard for an EDA (Environ- 
mental Damage Assessment). It is surprising that the 
exact number on “how many seabirds are really 
oiled” is hard to get and not available with high accu- 
racy; accurate numbers seem not to play a role in the 
legal decision and discussion even! Perhaps court 
fines should consider a price per oiled seabird, and 
thus could change the current dilemma?! 

This document reports that approximately 40 mil- 
lion pelagic seabirds reside during the year on the 
Grand Banks off Newfoundland. However, some of 
the presented numbers are puzzling and might cause 


Book REVIEWS 30 


Oo 


confusion for the informed Naturalist. It was report- 
ed earlier that over 200 000 Thick-billed murres are 
killed annually during the Murre hunt off Newfound- 
land. Now, chronical oil pollution is even added, but 
breeding Thick-billed Murres in the Canadian Arctic 
— the seabird species believed to be affected the most 
by chronic oil pollution — does not show a signifi- 
cantly declining population trend at all. Are Canadi- 
an seabirds sensitive indicators of the marine envi- 
ronment ? Or are birds from other areas in the world 
and being present in Canadian waters, such as Green- 
land’s Thick-billed Murres and Manx Shearwaters 
from England (both populations are known to be 
declining), better indicators ? More research is 
required. Some other confusion might arise from the 
presented population numbers of wintering Eastern 
Harlequin Ducks, and that no direct relationship is 
known to exist between the amount of oil spilled and 
the numbers of seabirds killed. For my taste, some 
key references such as J. Burger’s 1997 book on “Oil 
Spills” would have been a great addition. Of interest 
might also be the seabird oil pollution work in 
British Columbia by A. Burger, the Festucca Oil 
Spill Trust Fund and the Provincial Government’s 
work. Globally speaking, it might be interesting for 
the reader to learn how Norway, a country with major 
offshore oil resources and with a very long coastline 
and huge seabird resources, deals with chronic oil 
pollution! Perhaps it would also be informative to 
have a list of all known oil vessel accidents in Cana- 
dian waters. 

However, this informative report provides many 
important details and baseline information on the 
sligthly overlooked but very relevant chronic oil pol- 
lution topic in the offshore waters of Eastern Canada. 
It focuses on seabirds; but many other species and 
the entire ecosystem suffer from oil pollution, too. 
“Chronic oil pollution is an international problem 
whose solution requires national and international 
effort”. 

FALK HUETTMANN 


Biology and Wildlife Department, University of Alaska, 
Fairbanks, Alaska 99777-7000 USA 


Spiders of Australia: An Introduction to their Classification, Biology and Distribution 


By T. J. Hawkeswood, 2003. Pensoft Publishers, Geo Milev 
Street 13a, 1111 Sofia, Bulgaria. 264 pages. EURO 19.95 
paper, 34.95 cloth 


The past few decades have seen the production of 
/a considerable number of landmark volumes on the 
| natural history of regional spider faunas. Volumes by 
Dippenaar-Schoeman and Jocqué (1997), Song et al. 
(1999), Ubick et al. (2005) among others have set a 
high standard for concise, useful, in-depth coverage 
of regional faunas. With this in mind, I readily agreed 
to review Spiders of Australia: an Introduction to their 
Classification, Biology and Distribution. 


I tried, I really did try, to find good things to say 
about this book. Certainly the dust cover of this vol- 
ume, with its border of 24 colour images of various 
Australian spiders surrounding an anthropomorphic 
photograph of the front end of an immature male 
deinopid spider (looking as charmingly pugilistic as 
only an immature male can), promises an interesting 
and engaging piece of work. Unfortunately, as they say, 
you can’t judge a book by its cover. 

The content of Spiders of Australia falls far short of 
fulfilling the promise of either the cover or title. Inter- 
esting observations on the natural history of a variety 


304 


of relatively common and fairly well known spiders 
from Oz are marred by errors of fact and/or presenta- 
tion on nearly every page. One is left with the impres- 
sion that the well-meaning author has little profes- 
sional knowledge of spiders and that the text never 
received professional content or copy editing. 

This volume commences with a brief preface fol- 
lowed by several short introductory chapters covering 
spider morphology, natural history, classification and 
other general topics. Anyone with more than basic knowl- 
edge of spiders will find much to criticize or question 
within these pages. For instance, we are presented with 
the following dubious “facts.” 

e Spiders are important in controlling mosquito pop- 
ulations. In reality, fish, birds and various other 
organisms are vastly more important for mosquito 
control than are spiders. 

¢ There are about 20 000 and 1800 spider species in 
the world and Australia respectively. The actual fig- 
ures are closer to 40 000 (Platnick 2006) and 3300 
(R. J. Raven, Queensland Museum, personal com- 
munications). 

e Male and female spider genitalia fit together like a 
“lock and key mechanism.” Wrong! Spider geni- 
talia are one of the best known examples of rapid 
evolution likely resulting from sexual selection by 
female choice (Eberhard 1985). 

¢ Palps of male spiders are “hollowed out” to hold 
sperm. An insult to male spiders everywhere! Mature 
male spiders are unique in (and defined by) their pos- 
session of palps bizarrely modified into complex 
sperm storage and transfer organs. 

¢ A considerable number of Australian spiders pose 
‘a serious threat to humans in Australia.” Wrong! 
Among Australian spiders, only widows (Latrodec- 
tus hasselti) and a couple of funnel-web spiders 
(Atrax and Hadronyche spp.) pose any threat and 
true bites from these spiders are exceedingly rare 
(Isbister 2004, Isbister and Gray 2002). Australians 
are better off spending their paranoia time worry- 
ing about being hit by lightning. 

Things don’t get much better in the subsequent 
chapter (“Species described in this book”) which forms 
the bulk of the text. Approximately 125 pages discuss 
141 species in 29 families (or about 4 and a little more 
than 30% of the currently recognized species and 
families represented in Oz). Each family section lists 
the number of Australian and world species in the 
family and describes the general appearance, life his- 
tory, and behaviour of one or more exemplar species. 
Unfortunately, the families are organized in a confus- 
ing quasi-phylogenetic manner, the species statistics 
are often erroneous, the descriptive information is large- 
ly useless for identification purposes, other information 
is often misleading or erroneous, and the vast majority 
of the unique and truly wonderful Oz spider fauna is 
ignored. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


For instance, consider the author’s treatment of the 
family Pholcidae. Australian genera and species are 
listed as 9 and “about 12” and the cosmopolitan Phol- 
cus phalangioides is the token exemplar. The author 
references the most recent taxonomic work summa- 
rizing the Oz pholcid fauna (Huber 2001) but missed 
the fact that this work records 14 genera and nearly 
80 species (and still counting). Well, okay, I some- 
times have trouble with math, too. But in such a book 
I expect to be introduced to truly Australian pholcids 
instead of to an already well-known species found 
throughout the world. The section is further marred 
with factual and typographical errors. Similar prob- 
lems surface in the treatment of other families. As well, 
erroneous medical “information” is regularly trun- 
dled forward — e.g. under Lamponidae, almost entirely 
an Australian family, discussion of the life and times 
of Queenvic piccadilly would be vastly preferable to 
reiteration of the discredited medical mythology sur- 
rounding Lampona cylindrata. 

A short glossary of slightly less that 100 entries 
follows the “Species described ...” chapter. Explana- 
tions are generally clear but one wonders why such 
terms as anus, bark, and solitary warrant entries but 
mygalomorph does not. Nearly 30 pages of refer- 
ences conclude the text. There is an annotated section 
on general texts on Australian spiders and fairly com- 
plete and up-to-date scientific references are present- 
ed for each family. In spite of considerable repetition 
[e.g. Rainbow (1911) appears no less than 29 times] 
the references are easily the most useful part of the 
entire book. 

The book finishes with a set of 139 photographic 
plates and 27 paintings of the main species discussed 
in the text, arranged 3 to a page. Some of the photo- 
graphs are quite nice illustrations (e.g., plate 5 Deinopis 
subrufa, the cover boy mentioned earlier, and plate 16 
Lycosa bicolor, also featured on the cover as one of 
the border images). The paintings are without excep- 
tion unexceptional — two-dimensional, flatly coloured, 
and crudely rendered. 

In summary, if you are looking for an interesting 
decorative wall piece, frame the cover and recycle 
the rest (maybe keep the references section). If seri- 
ously curious about general spider biology and clas- 
sification, spend your money on Levi and Levi (1968) 
or Foelix (1996). For specific information on the spiders 
of Oz, buy Murphy and Murphy’s (2000) treatise on 


southeast Asian spiders — it does a much better job of | 


introducing the fauna than does Spiders of Australia 


References: 

Dippenaar-Schoeman, A. S., and R. Jocqué. 1997. African Spi- 
ders: an Identification Manual. Agricultural Research Council, 
Pretoria. 392 pages. 

Eberhard, W. G. 1985. Sexual Selection and Animal Genitalia. Har- 
vard University Press, Cambridge, 244 pages. 

Foelix, R. F. 1996. The Biology of Spiders, 2"¢ ed. Oxford University 
Press. 330 pages. 


2005 


Huber, B. A. 2001. The pholcids of Australia (Araneae; Pholcidae): 
taxonomy, biogeography, and relationships. Bulletin of the Ameri- 
can Museum of Natural History 260: 1-144. 

Isbister, G. K. 2004. Necrotic arachnidism: The mythology of a mod- 
ern plague. The Lancet 364: 549-553. 

Isbister, G. K., and M. R. Gray. 2002. A prospective study of 750 def- 
inite spider bites, with expert spider identification. Queensland 
Journal of Medicine 95: 723-731. 

Levi, H. W., and L. R. Levi. 1968. A guide to spiders and their kin. 
Golden Press, New York. 160 pages. 

Murphy, F., and J. Murphy. 2000. An introduction to the spiders of 
South East Asia with notes on all the genera. Malaysian Nature 
Society, Kuala Lumpur. 689 pages. 


BOOK REVIEWS 


Platnick, N. I. 2006. The world spider catalog. Version 6.5. The 
American Museum of Natural History. http://research.amnh.org/ 
entomology/spiders/catalog/index.html 

Rainbow, W. J. 1911. A census of Australian Araneidae. Records of 
the Australian Museum 9: 107-391. 

Song Daxiang, Zhu Mingsheng, and Chen Jun. 1999. The spiders 
of China. Hebei Science and Technology, Shijiazhuang. 640 pages. 

Ubick, D., P. Paquin, P. E. Cushing, and V. D. Roth (Editors). 2005. 
Spiders of North America: an Identification Manual. American 
Arachnological Society. 377 pages. 

ROBB BENNETT 


Seed Pest Management Officer, BC Ministry of Forests, 7380 
Puckle Road, Saanichton British Columbia V8M 1W4 
Canada 


Annotated Bibliography of Quaternary Vertebrates of Northern North America With 


Radiocarbon Dates 


Edited by C. R. Harington. 2005. University of Toronto Press, 
10 St. Mary Street Suite 700, Toronto, Ontario MST 1R5 
Canada. 539 pages. $150.00 CDN. 


Bibliographies have proven an essential tool in any 
historical based research, yet are often underrated. In 
paleontology, specifically vertebrate paleontology, many 
have relied upon Bibliography of Fossil Vertebrates 
Exclusive of North America 1509-1927 (Romer et al., 
1962), and Bibliography of Fossil Vertebrates (BFV) 
(produced by the Society of Vertebrate Paleontol- 
ogy), and their earlier versions. The BFV, however, is 
no longer updated. Taxon-oriented bibliographies are 
also produced, like Crossman and Casselman’s (1987) 
annotated bibliography of Esox lucius; and theme ori- 
ented bibliographies like Tokaryk et al. (1992) anno- 
tated bibliography of the Cretaceous-Tertiary extinction 
event. Without continued maintenance, these, however, 
are quickly dated. 

Harington’s Annotated Bibliography of Quaternary 
Vertebrates of Northern North America — with Radio- 
carbon Dates is a recent contribution to the stacks of 
paleontological resources. This volume contains 1347 
citations (in 328 pages) from 1748 to 2000, containing 
descriptions or notices of fauna from 2 million to 5000 
years ago. The region is inclusive of Alaska, Greenland, 
and Canada. 

The annotation is of sufficient depth when warrant- 
ed and the reader will note the brevity given to many 
of the pre-20" century articles as these themselves lack 
sufficient depth. The indexing, always a vital tool in 


White as a Ghost: Winter Ticks and Moose 


By Bill Samuel. 2004. Federation of Alberta Naturalists, 
11759 — Groat Road, Edmonton, Alberta, TSM 3K6 Canada. 
100 pages. $ 24.95. 


Parasites are a fascinating study. They are able to 
adapt and evolve in order to survive in or on their hosts, 
but usually will not kill their hosts. Books about par- 
asites are not often found in a public library, but White 
as a Ghost would be a good addition. It has a large for- 
mat, 28 cm x 23 cm, with excellent photographs illus- 


bibliographic construction, is subdivided into five sec- 
tions: scientific names; common names; localities and 
stratigraphic terms; personal names and institutions; 
and of a general index. This latter section, always con- 
strained by the subjective nature of its composer, can 
be relied upon for consistency simply for the fact of 

Harington’s long time standing in Quaternary pale- 

ontological research, which is beyond reproach. 

The facet that will make Harington’s work not only 
stand out but retain a longer shelf life 1s the 138 addi- 
tional pages devoted to radiocarbon dates associated 
with taxa. This extended table, all the more important 
in Quaternary paleontology than in any other paleon- 
tological subdivision, includes institutions responsible 
for the results, and published references. Collectively, 
this volume will provide a long lasting reference for 
students and professionals of Quaternary life in North 
America. 

References: 

Crossman, E. J., and J. M. Casselman. 1987. An Annotated Bibli- 
ography of the Pike, Esox lucius (Osteichthyes: Salmoniformes). 
Royal Ontario Museum, Toronto, 386 pages. 

Romer, A. S., Nelda E. Wright, Tilly Edinger, and Richard Van 
Frank. 1962. Bibliography of Fossil Vertebrates Exclusive of North 
America, 1509-1927. The Geological Society of America, Memoir 
87, 1544 pages. 

Tokaryk, T. T., J. E. Storer, and E. M. V. Nambudiri. 1992. Selected 
Bibliography of the Cretaceous-Tertiary Boundary Event, Through 


1989. Natural History Contributions, Number 11. Saskatchewan 
Museum of Natural History, Regina, 140 pages. 


TIM TOKARYK 
Box 163, Eastend, Saskatchewan SON OTO Canada 


trating the text, but is not a coffee table book. It is 
intended to give trappers, Fish and Wildlife officers, 
hunters, farmers and biologists the knowledge they need 
to understand and recognize the life cycle of the tick 
which causes Ghost Moose: Dermacentor albipictus. 
Wilderness campers, more than the general public, are 
most likely to come across Ghost Moose. 

Dr. Samuel is a parasitologist who studies para- 
sites of deer, Elk and Moose and in particular the tick 
which causes Ghost Moose. The tick is widespread 


306 


and is found throughout the North American ranges 
of deer, Elk and Moose except the far north. Deer and 
Elk seem able to co-exist with this parasite, and they 
remove most of the ticks by efficient grooming before 
they can do much harm. But Moose are less capable 
of combating an infestation. The eggs of D. albipictus 
are laid on vegetation in the spring, and from Sep- 
tember to November the hatched larvae climb up the 
vegetation and attach themselves to a passing animal. 
After November the larvae left on the ground die. Once 
attached to Moose, larvae feed on Moose blood before 
becoming nymphs; the nymphs are dormant until Feb- 
ruary, then moult and become adult ticks. All three 
stages depend on blood to survive, but it is in March 
and April that their feeding causes the Moose to be- 
come so weakened by loss of blood that some can die. 
The tick bites can also introduce other pathogens into 
the host so that there may be multiple causes for death. 
After May, the female ticks drop off the host, lay their 
eggs in sheltered places and the cycle starts again. 
The irritation of the bites make the Moose groom in- 
cessantly, mostly with its tongue, which breaks off the 
ends of the coat hair leaving only the short undercoat 
exposed. This is grey-white and gives the Moose the 
typical “ghost” appearance. Licking can also break the 
skin, allowing easier access for the ticks. A serious 
infestation will create large open areas of bleeding skin 
and consequent debilitation. 

Moose populations in Canada are stable, but when 
a local population increases, the young in particular 
start the winter under-nourished and more suscepti- 
ble to disease. Then there can be a serious die-off in 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


March and April due to tick infestation. There were 
major die-offs of up to 50% of the population in some 
areas in 1991 (Minnesota), 1999 (six provinces, British 
Columbia the worst), and 2002 (eastern provinces and 
the prairies). Some dead Moose have carried more 
than 500 000 ticks. Effective prevention is difficult but 
Dr. Samuel suggests that a possible solution would 
be to cull Moose where there is a population build up, 
thus reducing the demand on the available food sources. 
Experiments have shown that Moose can detect and 
avoid tick-infested vegetation but if food is scarce, 
they will eat it. 

There are chapters on the life cycle of the tick, how 
they are adapted to attack Moose, their invasive char- 
acteristics and behavioural strategies used by Moose 
to evade the ticks. The book is attractively illustrated 
and for light relief, there are good puns on the word 
“tick”: e.g., characteris-ticks, and a poem or two. 
Knowing more about the life cycle of the tick and its 
serious effects on Moose populations should be of 
value to anyone travelling or working in the areas 
where moose are found. 

The Federation of Alberta Naturalists has published 
several natural history books, and their Atlas of Breed- 
ing Birds of Alberta is a best seller. They commis- 
sioned Dr. Samuel to write Ghost Moose which is an 
interesting and serious book about a wildlife disease. 
It makes absorbing reading and describes an aspect 
of wildlife we seldom hear about. 


JANE ATKINSON 
255 Malcolm Circle, Dorval, Quebec H9S 1T6 Canada 


Lewis and Clark on the Great Plains: A Natural History 


By Paul A. Johnsgard. Bison Books, University of Nebraska 
Press, 1111 Lincoln Mall, Lincoln, Nebraska 68588-0630 
USA. 143 pages, $18.28. 


One of the most profound expeditions in American 
history and culture was of the Lewis and Clark adven- 
tures, 1803-1806. Led by Meriwether Lewis (1774- 
1809) and William Clark (1770-1838), the expedition 
was one of the first systematic surveys of the natural 
resources and natural history of the American west. 
And as such, the published record since the early 19" 
century pertaining to these treks is voluminous. From 
narratives and journals (e.g., Moulton 1983-2002), 
their place in history, and their experiences with natu- 
ral history (Cutright 1969), all raise Lewis and Clark 
to American cultural icons. 

Johnsgard’s Lewis and Clark on the Great Plains 
adds to this list but should not be considered a signif- 
icant contribution to the histrionics of the expedition 
and what it meant. The author’s contribution is more 
in line with a combination field guide / history lesson 
of the wildlife encountered by Lewis and Clark’s group. 
Following a brief introduction, the book is chaptered 
by contemporary States with regional maps outlining 
expeditionary routes. This template is essential in un- 


derstanding the chronology as the expedition at times 
retraced their steps in a single season, if not over the 
course of the expedition’s history. 

The bulk of the content is a listing of the flora and 
fauna (complimented with scientific and common 
names). In user friendly manner Johnsgard provides a 
concise synopsis of the taxa followed by shorter nota- 
tions as to the encounters with the expeditionary force. 
The Western Hognose Snake (Heterodon nasicus) for 
example, was likely found and described on July 23, 
1805 near Townsend, Montana, prior to the formal 
erection of the species in 1852 (Baird and Girard 1852). 
Based on his description, “Lewis should be credited” 
Johnsgard contends, “with the discovery of the spe- 
cies” (page 97). 

Visual support for the descriptions come from the 
authors own line drawings, 39 in all. Simple in vision 
yet detailed, collectively with the body of text, make 
this little volume an added historical perspective to 
viewing nature as it once was, 200 years ago. 


References: 

Baird, S. F., and C. F. Girard. 1852. Reptiles. Pages 336-353 in | 
Exploration and Survey of the Valley of the Great Salt Lake of | 
Utah, including a reconnaissance of a new rout through the Rocky | 


2005 


Mountains. Edited by H. Stansbury. Lippencott, Grambo, and 
Company, Philadelphia, 487 pages. 

Moulton, G. E. Editor. 1983-2001. The Journals of Lewis and Clark 
Expedition. 13 volumes. University of Nebraska Press, Lincoln. 


Waterfowl of Eastern North America 


By Chris G. Earley. 2005. Firefly Books: Buffalo, New York; 
and 3680 Victoria Park Avenue, Toronto, Ontario M2H 3K1 
Canada. 158 pages. $19.95 Paper. 


This attractive bird identification book is clearly 
aimed at the beginner birder. It is the fourth in a series 
of similar books by the same author and publisher. 
Earley properly cautions new birders about hasty and 
uncertain judgments in the often perplexing task of bird 
identification. 

A cursory review of this handsomely printed, full- 
color book, measuring 54" (14.0 cm) by 84" (21.6 cm) 
by 4" (1.3 cm), makes a favorable first impression. In 
particular, the photographs of individual birds, usually 
in nuptial plumage, on the water and flying, and when 
used in comparison with look-alike birds, are excel- 
lent. Two pages are usually devoted to each species, 
including photographs, a small range map, and brief 
text mostly descriptive of non-breeding season plum- 
ages. Two to four sentences tell a bit about each species 
and a sentence or two under “Nature Notes” provides 
additional eclectic, often trivial information. 

Unfortunately, my initial favorable impressions 
soon changed. First, the title is a misnomer. A substan- 
tial number of the species included are not waterfowl 
as defined by the AOU checklist, which the author 
references. Rather, the author re-defines waterfowl to 
include other “ducklike birds”, such as some species 
of loons, grebes, pelicans, cormorants, rails, and galli- 
nules. Attention to these species comprises about a 
quarter of the pages accorded the true waterfowl. 

North America and Eastern North America are not 
defined. By AOU definition the mainland of North 
America extends south through Panama, including 
associated islands. Consequently, some native North 
American species are missing. On the other hand, sev- 
eral exotic or species of rare or irregular occurrence are 
addressed (e.g., Mute Swan, Barnacle Goose, Garganey, 
Tufted Duck, and Smew). Oddly, four full pages are 
devoted to each of three species rarely (unlikely ever 
to be) seen by many birders (the King and Common 
eiders and the Long-tailed Duck); but the commonest 
North American duck, the Mallard, receives only two 
pages. 

Other problems arise. The range maps are too small, 
especially for coastal species when the pale yellow 
representing winter range is cast against the light gray 
background. The map depicting the wintering range 
of the Blue-winged Teal (page 47) as including north- 
eastern Argentina, Paraguay, Uraguay and extreme 
southern Brazil is incorrect. Upon close examination, 


Book REVIEWS 


307 


Cutright, P. R. 1969 [1989]. Lewis and Clark: Pioneering Natural- 
ists. [Reprint]. University of Nebraska Press, Lincoln. 


TIM TOKARYK 
Box 163, Eastend, Saskatchewan SON OTO Canada 


the unnatural, vertically-flying shoveler (page 9) is 
certainly a cropped view of the horizontally-flying bird 
depicted on page 14 with the photo reversed and rotat- 
ed 90 degrees. The photograph of a flying male Com- 
mon Merganser (page 147) is mis-labeled as a Hooded 
Merganser. 

The two-page chart titled “Seasonal Status of Water- 
fowl” (pages 16-17) is based solely on observations at 
Point Pelee National Park [Ontario]. While perhaps 
useful in showing relative species abundance and tem- 
poral distribution in the Great Lakes region, it has 
questionable relevance elsewhere. 

“What can I do to help waterfowl?” (pages 130- 
131). is largely about nesting boxes. Only a half-dozen 
or so waterfowl utilize such structures. Nothing is said 
about North American waterfowl being protected by 
international treaties, federal laws, and laws and reg- 
ulations of the various states and provinces. 

The list of references (pages 154-155) is peculiar. 
Some entries are outdated (e.g., Studer, 1881) or are 
primarily of works of art (e.g., Brasher 1962, Lans- 
downe 1980). While the definitive AOU The Birds of 
North America monographs (Poole and Gill, eds.) are 
noted, their scientific excellence, content, and format 
are undescribed and their general unavailability, except 
in larger libraries, is unmentioned. 

A number of important waterfowl references are 
overlooked (e.g., Delacour’s four-volume The Water- 
fowl of the World; Palmer’s Handbook of North Amer- 
ican Birds, Volumes 2 and 3, Waterfowl; Volume 1 
(Ostrich to Ducks) of the magnificent Handbook of 
the Birds of the World, edited by del Hoyo, Elliott, 
and Sargatal; and even the early but enduring John C. 
Phillips’ four-volume opus, A Natural History of the 
Ducks. Despite the attention given wood ducks and 
nesting boxes, Frank Bellrose’s splendid monograph 
(Ecology and Management of the Wood Duck) 1s like- 
wise missing. All of these basic references could easily 
have been accommodated in the large blank space on 
page 155. 

Much of what the novice birder learns in this book 
will have to be forgotten should his early interest in 
birding lead him forward. Perhaps the concerns and 
errors noted above, and others unexpressed, will be 
corrected in a reprinting. In the meantime one would 
be wiser to spend a few more dollars for one of several 
time-tested, authoritative field guides listed among 
the author’s references. 


HENRY M. REEVES 
22250 Boulder Crest Lane SE, Amity, Oregon 97101 USA 


308 


Wolves: Behavior, Ecology, and Conservation 


Edited by D. Mech and L. Boitani. 2003. University of Chica- 
go Press, 5801 Ellis Avenue, Chicago, Illinois, 60637 USA. 
xvii + 448 pages, U.S. $49 


In 1961, wolf biologist Douglas Pimlott wrote: “The 
wolf poses one of the most important conservation 
questions of our time. Will the species still exist when 
the twentieth century passes into history?” 

Pimlott, if he were alive today, would be amazed 
and heartened by Wolves: Behavior, Ecology, and Con- 
servation by L. D. Mech and L. Boitani because not 
only does it describe evidence of a “turn around” for 
the species that occurred in recent decades, but pro- 
vides an encyclopaedia of scientific information about 
the wolf that has come from a remarkable amount of 
research over the past 40 years. 

This book replaces one by David Mech written in 
1970. However, this “update” is a much expanded, 
22-authored, 448 page tome that provides one of the 
most extensive descriptions of the ecology of any 
mammal species in the world. The Table of Contents 
provides evidence of the book’s depth, with chapters 
on: wolf social ecology; behaviour; communication; 
wolf-prey relations; wolf physiology; genetics; evo- 
lution and taxonomy; interactions with non-prey; res- 
toration of the red wolf; wolves and humans; wolf con- 
servation and recovery. 

This is a book that you should read in selected chap- 
ters, rather than from cover to cover. There is too much 
detail to absorb, and although mostly written clearly, 
the style is fully referenced science. The book is well 
indexed, and brings together research conclusions on 
any conceivable topic related to wolves, very useful 
either for the interested person or the biologist. 

The intent of the book, according to the editors, is 
to counter the “myth and legend, forklore and fairy tale” 
that has, and continues, to surround the species, by 
presenting a scientific view of the animal. The under- 
lying assumption is that this scientific understanding 
will result in support for wise management of the ex- 
panded wolf populations that now exist in many places 
in the world. A more poetic reason for the book is 
given by physiologist Terry Kreeger: “Physiologically, 
we know a great deal about the wolf, although we still 
have much to learn. But why should we continue to 
study the wolf? Some people curse the animal; others 
deify it. As scientists study it, we may be able to blunt 
these extremes and place the wolf in proper perspec- 
tive. Wolves tend to roughen the edges of a world being 
smoothed by human hands. For many of us, that is 


BOTANY 


Rendezvous with the Wild: The Boreal Forest 


Edited by James Raffan, 2004. The Boston Mills Press, 132 
Main Street, Erin, Ontario NOB 1T0O Canada. 192 pages, 
$49.95 Cloth. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


good reason to learn what we can about them, inside 
and out, and certainly good reason to work for their 
conservation.” 

Some chapters are difficult, particularly those ad- 
dressing wolf taxonomy and genetics, because the data 
from research are themselves confusing, contradictory 
or only tentative. Both fields have been fraught with 
““re-interpretations,” sometimes by the same researchers, 
and consensus on what constitutes adequate evidence 
is unclear. For example, in the genetics chapter is a 
statement that parent-offspring relationships can be 
determined by examining nuclear DNA at as little as 
10 microsatellite loci, whereas work done in associa- 
tion with our Algonquin wolf studies showed that as 
many as 15 loci were needed to avoid mistakes. These 
chapters leave their respective topics in chaos; hope- 
fully, analytical methods soon will improve. 

All other chapters are more readable and, because 
of the wealth of data, lead to more intruiging descrip- 
tions of the lives of wolves. Portrayed here are images 
of the wolf as a highly adaptable and intelligent species, 
one with a set of biological limits and norms, but 
with the flexibility to exercise a great deal of individual 
choice, the key to its success. 

Human-wolf relationships are chronicled through 
the ages, right up to modern attitudes and their con- 
sequence for the future of the species. In a final chap- 
ter, Mech and Boitani reflect on the need to shift our 
perspectives on how to manage wolves from one of past 
“trench warfare” between people with different atti- 
tudes, to some new, more moderate paradigm that 
accepts the wolf with human imposed limits on pop- 
ulation size, particularly in human-altered environ- 
ments where it has been shown capable of surviving. 
Missing, however, is recognition of the importance of 
maintaining at least some areas as a crucible of natu- 
ral selective forces surrounding the species — the very 
forces out of which the species evolved — rather than 
being content with the imposition of human modified 
environments and human control. There is still more 
to achieve before we congratulate ourselves in saving 
the real “wild” wolf in real intact wilderness. 


Literature Cited 
Pimlott, D. H. 1961. Wolf control in Canada. Canadian Audubon 
Magazine, November-December, 2-9. 


JOHN B. THEBERGE 


RR3, Site 25, Compartment 82, Oliver, British Columbia 
VOH 1TO Canada 


Named after the Greek god of the northwind, Boreas, 
the boreal forests of the world carpet the northern cir- 
cumpolar reaches. St retching across Canada from New- 


2005 


foundland to the Yukon, the boreal forest reaches into 
Alaska, through the vastness of Russia and into the 
Nordic countries of Scandinavia. Forests of coniferous 
black spruce, white spruce, balsam fir, jack pine, and 
tamarack, interspersed with deciduous white birch, 
aspens, willows and alders dominate the boreal scene. 

Canada’s boreal forests are of national, and indeed, 
global significance. Approximately 40% of the globe’s 
boreal forests lie within Canada’s boundaries. Fifty- 
eight percent of Canada’s landmass is boreal forest 
that includes over 90% of the country’s remaining 
large, intact forest landscapes or 25% of the globe’s 
intact forests. Ecological values include prime habi- 
tat for many species of wildlife (including 75 percent 
of the continent’s waterfowl), vast areas of lakes, rivers 
and wetlands and globally significant storage of carbon. 

Canadian forests, especially the boreal forest, have 
long played a key role in the national economy. In 
2003 alone, forest products contributed almost $30 
billion to Canada’s $46 billion trade balance. Canada 
is the world’s second largest producer of wood pulp 
and the world’s largest producer of newsprint. Direct 
forest industry employment totaled 376 300 workers 
for 2003. Much of this economic activity is directly 
related to the boreal forest that acts as the economic 
foundation for many communities across the country. 

Given the economic and ecological significance of 
Canada’s boreal forest, it is small wonder that people 
are becoming increasingly concerned about the long- 
term sustainability of this continental biome. Rendez- 
vous with the Wild is the latest in a series of books, 
articles and media features on the future of the boreal 
forest. It tells the story of the Boreal Rendezvous, a 
series of canoe trips taken in the summer of 2003 on 
ten Canadian boreal rivers from the Wind River in 
the Yukon to the Moisie in Quebec. 

The canoe trips were a vision of the Canadian Parks 
and Wilderness Society (CPAWS) in collaboration 
with the David Suzuki Foundation, the Canadian 
Boreal Initiative, and Mountain Equipment Co-op. 
They sought ways that would help shift our relation- 
ship with the boreal forest from one of accelerating 
large-scale, industrial fragmentation and transforma- 
tion, to one that focused more on boreal forest con- 
servation and viable, sustainable development. 

Edited by the well-known Canadian author and avid 
canoeist James Raffan, Rendezvous with the Wild is 
an enticing collage of photography, art, journal entries, 
essays, poems, musings and prayers from many of 
the canoe trip participants. The variety of entries is 
marked by the diversity of contributors. Over 70 peo- 


ENVIRONMENT 


BOOK REVIEWS 


309 


ple including Native elders, conservationists, television 
celebrities, scientists, photographers, poets, academics, 
canoe builders, and musicians contributed their voice 
and creativity to this marvelous tribute to the mystery 
and attraction of the boreal forest. 

As Raffan notes in his opening essay, this 1s not a 
book about the boreal forest, but rather a book in res- 
ponse to the boreal forest. The canoe trips and canoe- 
ing act as constant themes weaving their way through 
the rich and varied fare. The photography is splendid 
and the book’s layout a delight to the senses. The 
reader is carried along the current of the river with 
boreal vistas provided by the many witnesses offered 
by the book. Of particular strength throughout the 
book is the vision and witness of the First Nations to 
their boreal forest home. 

The French philosopher Blais Pascal once remarked 
that the human heart has reasons of which the mind 
knows little. In boreal forest conservation issues, the 
environmental community often takes refuge in the 
technical, scientific dimensions of any particular bore- 
al forest issue. This is essential and necessary. How- 
ever, sole attention to the technical dimension fails to 
tap the depths of energy that can be attributed to the 
multidimensional human experience of the boreal for- 
est. This human “emotional” experience is often dis- 
missed as simply a “subjective,” private experience 
that cannot be accepted on par with so-called “objec- 
tive” scientific knowledge of any particular issue. Ren- 
dezvous with the Wild dispels such dualistic thinking 
and attempt to legitimize the direct human experience 
of the boreal forest as a powerful force that may ener- 
gize action on behalf of forest conservation. 

Rendezvous with the Wild begins and ends with a 
prayer by William Commanda, an Algonquin elder 
from Maniwaki and honorary elder of the CPAWS 
Boreal Program. The book is thus bounded by the 
spiritual, by due attention to the human experience of 
the boreal forest. You will have to look elsewhere for 
material on the boreal forest, on its ecology, on the im- 
pact of industrial activity, or the development of boreal 
forest policy. Rendezvous with the Wild attends to other 
data, to the inner data of human consciousness vis-a- 
vis the boreal forest. I have no doubt that if such data 
is not seriously considered, then conservation and 
sustainable development of Canada’s boreal forests 
will remain a dream — forever. 


JOHN MCCARTHY 


Holy Rosary Parish, 175 Emma Street, Guelph, Ontario 
NIE 1V6; jmccarthy @jesuits.ca 


The Earth’s Blanket: Traditional Teachings for Sustainable Living 


By Nancy J. Turner. 2005. University of Washington Press, 
P.O. Box 50096, Seattle, Washington 98145-5096 USA. 
298 pages, U.S. $29.95. 


Nancy Turner’s book, The Earth’s Blanket is a thor- 
ough treatise on indigenous peoples’ relationships 
with the environment, and has as its goal to demon- 


310 


strate and better understand alternative ways of view- 
ing the world. Underlying the theme is the concern 
that the rich environmental knowledge that sustains 
the earth’s ecosystems 1s being lost at a time of great 
environmental devastation. It is well researched, well 
supported with references and source notes, and cites 
numerous examples to substantiate all claims. 

The central thesis revolves around the concept of 
the earth’s blanket, which is a metaphor used by the 
Nlaka’pmx of the southern interior of BC, to describe 
the plants that cover the earth; if removed will cause 
the earth to be “angry” and to “weep”. Turner weaves 
the theme of the reciprocal relationship that humans 
have with their environments throughout the eight 
chapters. The reader moves through discussions of 
wealth and value in a changing world, a kincentric 
approach to nature, honouring nature through cere- 
mony and ritual, to land stewardship, all richly illus- 
trated with stories of First peoples and historical ac- 
counts (from the 1800s onwards from the journals of 
Simon Fraser) of Indian agents, anthropologists and 
early European travelers. 

An ethnobotanist for over 30 years and as a non- 
indigenous academic, Turner skillfully combines the 
perspectives of indigenous peoples: chiefs, friends, 
Elders, ethnobotanists, in several communities in Bri- 
tish Columbia, with some examples from indigenous 
communities in other parts of the world (i.e. Sierra 
Terrahumara, in Mexico). Turner has fluency with First 
Nations peoples and issues in British Columbia and 
through developing trust and friendship, has become 
close to special and profound relationships with earths’ 
offerings. Turner provides several traditional stories, 
ceremonies and rituals that are “situated”, connected 
to the history and geography of the region and thus 
connects people to place. Stories also demonstrate how 
knowledge and understanding of the environment is 
relational — that the plants, animals and other features 
are imbued with human qualities, so that humans are 
not viewed as separate and outside ecosystems. 

Several important issues emerge throughout the 
book that tie together the loss of cultures and envi- 
ronmental destruction: the importance of language is 
mentioned several times, in that language is a reser- 
voir of traditional knowledge, culture and connections 
to the landscape. Loss of language severs that con- 
nection. The consequences of environmental destruc- 
tion are poignantly described using several species, 
with salmon a recurring example throughout the book. 
Salmon is a strong marker of cultural identity for Bri- 
tish Columbia First Nations and is a thread through- 
out the book that weaves together origin stories with 
scientific understandings of salmon ecology. Turner 
provides details on how to prepare salmon and chroni- 
cles the demise of salmon stock and the effect this has 
on communities that depend on the catch. One Sec- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


wepemce story about salmon migration helps children 
learn that salmon are to be respected and admired. 
Other key species that are indicators of ecological 
decline are the Bitterroot (Lewisia rediviva) used by 
the Nlaka’pmx peoples, numerous edible berries and 
abalone. 

In Chapter 7, “Everything is One”, Turner ties togeth- 
er the several different themes that focus on human’s 
interdependence with the environment. The intercon- 
nections that are so vital are expressed in the story of 
the Xaxl’ep people who live at Fountain, a small set- 
tlement near the Fraser River, where “Everything in 
their territory is connected to them and if part of it is 
lost the Xaxl’ep lose part of themselves.” This makes 
the point that when habitat and resources are taken 
away, people lose the knowledge associated with that 
place and that resource. 

Towards the end of the book, Turner introduces 
scientific concepts such resilience, complex systems, 
adaptive management and ecosystem based manage- 
ment to bring together the multi-faceted human- 
ecosystem relationship and how we can live sustain- 
ably. There are stories of hope and ideas for a 
sustainable future. For example, the St6:16 people of 
the Fraser valley are revitalizing their language and 
cultural heritage after the drainage of the Sumas Lake 
in the 1920s and the subsequent loss of traditional 
resources, and means of transportation for the St6:16 
people. The Tmix” research project of the Nlaka’ pmx 
people encourages developers to integrate traditional 
knowledge with land management to foster a sustain- 
able land ethic. Turner suggests three criteria for pos- 
itive change and eight concepts for ecocultural restora- 
tion and urges us to continue to be optimistic despite 
the level of environmental destruction that we are wit- 
nessing globally. 

If there are any shortcomings to this book, they are 
few. At times, it is difficult to delineate the chapters 
as several of them overlap and appear repetitive. While 
Turner is meticulous in providing both scientific and 
indigenous/local language names for all species men- 
tioned, the scientific name is sometimes provided more 
than once, when once would have been sufficient. Also, 
Turner does not address changes in attitudes towards 
the environment by First Nations — there are often 
difficult trade offs to be made when economic bene- 
fits seem to outweigh traditional teachings of respect 
and environmental conservation. Attitude change occurs 
within communities as values shift — addressing what 
is to be done about the loss of sense of responsibility 
for stewardship is a difficult issue. In all, however, 
the book provides important lessons on First Nations 
stewardship and promises of positive change if we all 
just listen. 


ELLEN WOODLEY 
RR4, Fergus, Ontario NIM 2W5 Canada 


2005 


BOOK REVIEWS 


311 


Greenpeace: How a Group of Journalists, Ecologists and Visionaries Changed the World 


By R. Weyler. 2004. Raincoast Books, 9050 Shaughnessy 
Street, Vancouver, British Columbia, V6P 6ES Canada. 
574 pages. $25.95. 


Rex Weyler has released an honest, informative, 
and politically motivating history of the first nine 
years of the Greenpeace movement, charting its course 
from its inception in a Vancouver living room in 1970 
to its official internationalization in an Amsterdam 
pub in 1979. While others have written histories of 
this now widely-studied movement, Weyler’s is of a 
different sort: he is an insider to the group, having been 
a Greenpeace activist since 1974, and knows its key 
players better than most. He relates individual moti- 
vations and personalities, and shares both the bitter 
disputes between factions of the movement and the 
eco-political victories celebrated over pints of beer. 
Weyler divides his study into three sections: War and 
Peace, All Sentient Beings, and Political Ecology. 
Throughout, he relates the growth of the movement 
and its increasing lens of awareness in a style that is 
clear and captivating, teaching readers about the com- 
mitments of the movement and winning new environ- 
mental activists as he details the group’s adventures. 

Weyler begins by locating the global climate of 1970, 
as seen from the west coast of Canada. War in Vietnam 
continued to rage, and American draft dodgers were 
fleeing to Canada by the thousand. The public outcry 
against war built throughout the 1960s, and the ever- 
quickening pace of the international arms race daily 
increased the number of anti-war activists. Mean- 
while, the world’s superpowers invested billions to 
stay in military advance of competing nations. Nuclear 
bombs were dropped at a rate of one per week, often 
with total disregard to populations neighbouring test 
sites. These nuclear blasts were contaminating entire 
cities with fallout, and the toxic Strontium-90, a by- 
product of the nuclear tests, was spread globally 
through the atmosphere. Most frightening, perhaps, 
were the questions surrounding the bombs that con- 
tinued to be tested with ever-increasing rapidity. In the 
race to produce the biggest and strongest atomic 
bomb, each superpower allowed their testing to pro- 
ceed in advance of scientific answers as to the effects 
of the bomb. Physicists cast bets among themselves 
about whether the next bomb would ignite the atmos- 
phere. Civilians were intentionally uninformed of 
their proximity to the tests, to allow governments to 
test the effects of radiation on humanity. 

Into this scene of rising and potentially deadly 
conflict, Weyler introduces his Vancouverite “cast of 
characters,” including political journalist Bob Hunter, 
pacifist ex-soldier Ben Metcalfe, and Quaker social 
activists Dorothy and Irving Stowe. This team realized 
that nuclear testing and the threat of nuclear warfare 
promised to kill our planet. They combined their 
strengths to develop a movement committed to gain- 


ing the general public’s support for pacifism, with 
emphasis on ending atmospheric nuclear testing. The 
team shared an awareness of the power of the media, 
and while they understood the science-based argu- 
ments against the danger of nuclear testing, they real- 
ized that the public would be won not by numbers, 
but by images. 

Hunter called these images “mindbombs”: they were 
“simple images, delivered by the media, that would 
‘explode in people’s minds’ and create a new under- 
standing of the world” (73). The team realized that 
whoever has the best picture wins, and set out to use 
this knowledge to their advantage. As ecological and 
disarmament goals merged, the group found their iden- 
tity advocating not only peace, but an environmentally- 
aware peace, a green peace, and thus found their name. 

The newly formed Greenpeace realized that protest 
groups had been largely ignored in the past because 
they weren’t demanding to be seen. Greenpeace decid- 
ed to sail a boat into the middle of the next scheduled 
nuclear explosion, a United States test scheduled to 
take place in October of 1971 on Amchitka Island, a 
“registered National Wildlife Refuge” (55). By plac- 
ing themselves in the middle of the event, and ensur- 
ing that dramatic photographs and news stories were 
released to the media, Greenpeace guaranteed that their 
campaign could not be overlooked. Though this ini- 
tial campaign, the sailing of the Greenpeace J and II, 
did not prevent the bomb from being detonated (cre- 
ating “the largest human-made earth tremor in history,” 
(131) a 7.2 magnitude earthquake), the mission was 
nevertheless a success. They had informed politicians 
and the general public of the dangers of the bomb, and 
had made their case for the futility of the arms race. 
As protests erupted around the world, atmospheric 
nuclear testing became an embarrassment to the Unit- 
ed States government, and experiments soon moved 
underground. Other countries followed suit, after sim- 
ilar pressure and embarrassing attention from Green- 
peace groups. 

The strategy proved successful: Greenpeace had 
lodged itself between the aggressor and the resource 
they aimed to protect, and people took notice. The 
movement grew, and attracted new activists. With the 
anti-nuclear success, the group shifted its attention to 
environmental injustices. Biocracy, the right of each 
living thing to be respected, became the credo of the 
group, as reflected in the “Greenpeace ecology mani- 
festo,” titled the “Declaration of Interdependence” 
(393). Individual Greenpeace members maintain indi- 
vidual ecological vision, and while this varies from 
member to member, most seem to be deep ecologists, 
maintaining that the survival of each living thing is 
dependent on the ecosphere at large; no species, in- 
cluding humans, takes priority over another. Green- 
peace as a whole adamantly proclaims its “fundamen- 


al? 


tal values” as “peace, tolerance, bearing witness, 
ecology, innovative direct action, [and] non-violence” 
(489). 

One of the new attendees at Greenpeace meetings, 
Dr. Peter Spock, a psychologist who had come to real- 
ize the enormous intelligence and complex brain waves 
of whales, advocated that their protection from whal- 
ing fleets should be the focus of the next Greenpeace 
campaign. 

Fleets of whaling ships, including factory boats 
that would process the whales while still at sea, had 
hunted the world’s whales to dangerously low levels, 
pushing some species to extinction. The Greenpeace 
team decided to emulate their successful anti-nuclear 
tactics, and again place themselves between the aggres- 
sor and the victim: they will pilot zodiacs between the 
whaling ships and the whales. This proved an enor- 
mously dangerous task, as the whaling ships were 
equipped with canon-fired harpoons, and the whalers 
were often not persuaded by the activists’ presence to 
hold their fire. Nevertheless, the Greenpeace team time 
and again positioned themselves between the guns and 
the whales, all the while snapping pictures to send to 
worldwide media, communicating the gory horror of 
the whale hunt. As with the anti-nuclear warfare cam- 
paign, Weyler relates devastating statistics about the 
depletion of the whale population, simultaneously ex- 
plaining the need for change and campaigning further 
to his readers. 

Both the anti-nuclear and the anti-whaling campaigns 
had found success, but the split-focus had divided the 
group. Names, places, and projects proliferate in the 
latter section of the book, speaking to the new nature 
and diversity of Greenpeace. However, funds were 
limited, and members disagreed about priorities. Fur- 
ther dividing the group was a new interest in an anti- 
sealing campaign. Sealers off the coast of Newfound- 
land and Labrador slaughtered hundreds of thousands 
of white coat seal pups each season, a hunt that Green- 
peace claimed was rapidly and dangerously depleting 
the seal numbers. Greenpeace activist Paul Watson 
advocated spraying the pups with a harmless green dye, 
which would make them worthless to the fashion 
industry to which their pelts are usually sold. The team 
set out for Newfoundland, with this intent. They were 
not well received by Newfoundlanders, who viewed 
them as outsiders, without right to decide on New- 
foundlanders’ means of income. 

Realizing the support of Newfoundland was vital, 
Hunter agreed to abandon the green dye idea, in favour 
of direct confrontation with the sealing companies. 
The focus, Hunter maintained, was not the sealers 
themselves, but the foreign factory ships that employed 
the sealers for very little, and sold the seal pelts at a 
great profit. This focus seemed to appease the New- 
foundland sealers, who guardedly welcomed the 
Greenpeace activists. While Weyler makes mention 
of the realization that this is an economic as well as 
an environmental problem, and briefly alludes to a 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


campaign to replace a portion of the wages of sealers 
who would agree to give up their work, this vital 
aspect of the problem is not pursued. The anti-seal- 
ing campaign continued, but did not present an eco- 
nomic alternative for sealers. 

The group became increasingly divided as Watson, 
frustrated over the slow pace of success in Newfound- 
land, advocated for a more aggressive approach in 
confronting environmental abuse. Greenpeace, how- 
ever, remained staunchly committed to non-violence, 
and Watson left to form his own campaign, the Sea 
Shepherd Society. A faction of Greenpeace remained 
committed to the seal campaign, and there seemed now 
to be three separate Greenpeaces, each competing for 
funds, each demanding that their own campaign was 
most important. A suggestion arose from the group: 
“If we’re ecologists, then let’s rise above our particu- 
lar issues to see the bigger pattern” (231). Despite 
their vision of an ecologically-just society, it became 
clear that, amidst all of their funding problems and 
internal bickering, they were not promoting balance, 
nor were they celebrating diversity. As their debt mount- 
ed, they gave in to pressure from Greenpeace groups 
that had formed worldwide to give up Vancouver con- 
trol, and merge with newer collectives to become Green- 
peace International. Though the Greenpeace campaign 
lives on, its focus is no longer Canadian, and Weyler’s 
retelling ends here. 

The continued international presence suggests that 
this truly is “the movement the world needs” (135). 
Projects continue to proliferate: as www.Greenpeace.ca 
clearly evidences, “the ecological crisis seems to be 
expanding on an exponential trajectory” (352). It will 
be of great benefit to those motivated to participate in 
today’s Greenpeace campaigns to know something of 
the roots of the movement. 

Weyler makes only brief mention of a common crit- 
icism leveled at Greenpeace; the rising feminist move- 
ment called the Greenpeace Foundation sexist, and 
perhaps rightfully so. Feminists accused the largely- 
male collective of militaristic language and macho 
and aggressive tactics. Hunter suggested at the time 
that a forceful and aggressive strategy was necessary to 
ensure that their environmental concerns were heard. 
Weyler himself doesn’t take the claims to task: instead, 
he allows the history of the movement to speak for 
itself. Throughout this history, he highlights the con- 
tributions of women to the campaign, and follows the 
story of Susi Newborn’s efforts to launch the first 
Greenpeace UK boat, the Rainbow Warrior, on a huge- 
ly successful mission. The concerns of ecofeminists, 
however, seem to stand, even as Hunter’s motto “A 
flower is your brother” (150) changed to the more in- 
clusive “A flower is your brother and your sister” (489). 
Other criticisms, such as the now common concern 
that the boats Greenpeace uses for its eco-interventions 
are themselves far from eco-friendly, remain unad- 
dressed. But perhaps such silence is justified: Weyler 
doesn’t set out for himself the task of justifying Green- 


2005 


peace. Instead, he recounts the passions that formed 
the movement and kept it motivated, and the inner 
conflicts that forced its evolution. 

The history certainly has its shortcomings. A tire- 
some Lord of the Rings analogy runs throughout the 
574 pages of text, and the foreshadowing of future 
Greenpeace trouble is almost constant. Most distract- 
ing is Weyler’s tendency to tediously set a scene, detail- 
ing the room, the lighting, and even the contents of 
paintings hanging on the walls. But his goal is as much 
to convey the emotion as the historical record of these 
first nine years, and the details help to transport the 
reader to the time and place remembered. (Weyler’s 
elaborate scene-setting becomes slightly more under- 
standable once we learn that he is the group photog- 
rapher. ) 


BooK REVIEWS 


313 


Ultimately, Weyler’s passion and enthusiasm for the 
ideals of the Greenpeace movement are shared with 
the reader in a style that is politically, scientifically 
and historically informed, making his book the per- 
fect starting point for anyone who is looking for 
either a history of Greenpeace, or the inspiration to 
become politically and environmentally active. This 
history, I think, will make Greenpeace’rs proud, as it 
functions as a mindbomb: reading about Greenpeace’s 
commitments, their successes and failures, awakens 
an awareness of the potentials of eco-activism within 
each reader. You can put the book down, but you 
can’t stop thinking about it... 


ERICA KELLY 


University of Western Ontario, London, Ontairo N6A 5B7 
Canada 


The Last Great Sea: A Voyage Through the Human and Natural History of the North Pacific 


Ocean 


By T. Glavin. 2000. David Suzuki Foundation and Greystone 
Books Douglas & McIntyre Publishing Group, Vancouver/ 
Toronto, Canada. 244 pages, $34.95 hardcover. 


This book belongs in every conservationist’s book- 
shelf, to say the least. As D. Suzuki describes very con- 
vincingly in the foreword, the environment of today’s 
North Pacific is characterized by its loss of (fish) species 
and its wipe-out of protein assemblages. The collapse 
of Sockeye Salmon is only one of many sad examples, 
many more exist: Steller’s Sea Cow, Spectacled Cor- 
morant, Dwason’s Caribou (Queen Charlotte Islands), 
and even plant species like Tobacco (Queen Charlotte 
Islands). Other species like Walruses, Sea Otters and 
Fur Seals barely survived until now. 

The first chapter starts slow but allows a very solid 
overview about historical and archaeological facts. 
Already after Chapter 2, nobody can deny anymore 
the environmental disaster and mis-management of 
the North Pacific and coastal British Columbia. Nev- 
ertheless, the author convinces the reader that the 
North Pacific still is THE largest fish producer in the 
world. “As in aboriginal fisheries, mythology played 
a part in industrial fisheries management, especially 
the myth of a superabundant ocean and the all-pow- 
erful capability of science and technology to fix the 
messes made by hydroelectric dams, lousy forestry 
practices and overfishing”. The governmentally encour- 
aged Merganser Control and Bear Shooting Programs 
designed for the sake of Salmon Protection prove this 
citation very well. Galvin strongly eliminates all illu- 
sions on how to heal the problem of overfishing. For 
instance, he shows that S. Livingstone’s widely fol- 
lowed idea of Fish Hatcheries does not produce more 
salmon, but instead takes away funds and harms nat- 
ural salmon stocks since they simply replace the last 
remaining and struggling stocks with poorly adjusted 
new ones. Strong also are Gavin’s arguments against 
Salmon Farming; e.g., it contributes to the closure of 


marine fisheries for wild salmon, and it requires 3 kg 
of fish to produce | kg of salmon. 

Fisheries and the ecology of all major North Pacific 
fish species get well-covered in this book. Since the 
abundance of salmon shaped western North America, 
this topic receives major attention in the text. All 
Pacific salmon species are discussed: Chum, Sockeye, 
Pink, Coho, Steelhead, Masu and Amago. Of major 
interest 1s in this regard the scientific discussion around 
the taxonomy of salmon; e.g., Steelhead (classified until 
1980s as Trout). The author brilliantly points out the 
implications of the religious-based and somewhat 
outdated taxonomical system by Carl von Linné, and 
how this affects the species management by national 
governments (provincial and federal) on an interna- 
tional level even (Canada vs. USA). 

The backwardness and failure of fishery laws are 
shown by outlining that the first salmon-fishing regu- 
lations for the Fraser River was a simple word-for-word 
replication of fishing regulations on English Rivers. 
At that time, Canada’s external affairs jurisdiction was 
still controlled by the British, which affects the Cana- 
da-U.S. salmon treaty concluded 1930s and renewed 
in 1985. In addition, Galvin shows that Canadian and 
U.S. fishery scientists significantly differed in their 
stock assessment results for the same species in the 
same waters even; consequently, so did the manage- 
ment and political agendas. This is the classical pic- 
ture of “mixed-stock” fisheries, which also threatens 
small salmon runs. 

The author reports the incidental death of 50 000 
marine mammals and 500 000 seabirds due to driftnet 
fishery activities in the North Pacific; marine (plastic) 
pollution comes with it. Despite the well shown fail- 
ure of a European and Western approach dealing with 
the North Pacific fisheries, domestic Japanese and 
Native fisheries seemed to work well and be sustain- 
able. Galvin shows the magnitude of “pre-contact” fish- 


314 


ery for salmon by natives, which was even compara- 
ble with levels of commercial fisheries from this cen- 
tury. Some readers might find that the book slightly 
follows stereotypical views of the noble native. 

A very strong point in this book is how the North 
Pacific and its fauna is linked with the “hinterland”: 
Old-growth rainforest, landscape and Bald Eagles. 
This needs to be considered in the light that resident 
Killer Whales in British Columbia are among the most 
contaminated cetaceans of the world. 

A very complete picture of the North Pacific is por- 
trayed by fully considering the Russian influence and 
history. The book outlines well that Russian settlers 
did much better than the western type of colonization 
(a point that might be put in doubt for the Kodiak 
Islands at least). The Russian-American Company was 
much more relevant in the history of North Pacific 
settlements and explorations than the Hudson Bay 
Company (HBC). But nevertheless, as with the HBC, 
the Russian quest for the North Pacific had the same 
motivation: central European pelt resources were al- 
ready overhunted! 

Regarding the marine ecology of the North Pacific, 
the importance of the Aleutian low, Pacific currents, 
and El Nino are fully described. This ecosystem is 
driven by “regime changes”, which calls for a dynam- 
ic management. The author outlines this very well by 
presenting the ground-breaking work from Russian 
Scientist T. Baranov, but also from Bill Ricker “Rick- 
er curve” and others at the Pacific Biological Station, 
e.g., G. McFarlane and D. Beamish. A quote from the 
book says it all: Understanding catch statistics is like 
“reading a single faded and crumbling onionskin page 
from an early draft of Wagner’s Tannhaeuser, in a dimly 
lit room”. Another quote of the book and taken from 
the U.N. Code of Conduct for Responsible Fisheries 
states, in part, that “the absence of adequate scientific 
information should not be used as a reason for post- 
poning or failing to take conservation and manage- 
ment measures”. Well said. 

Galvin makes a strong case that ethnocentric ap- 
proaches for understanding and managing the North 
Pacific have failed, e.g. the Chinese might have been 
in North America much earlier than the Europeans. 
The book elaborates on the major question “who came 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


first” since it had such a major economical and political 
implication for European powers. A major conclusion 
from this book is that there never was such a thing like 
an Old World (Eurasian) and New World (Americas). 

The chapters on anthropology and human history 
of the North Pacific and how the Russians, Asians and 
Natives settled and explored the North Pacific are on 
the same level than high-caliber books as Guns, Germs 
and Steel by J. Diamond. Just to name some high- 
lights, Glavin mentions how natives grew Arrowhead 
and potatoes, he cites the work of the Russian Anthro- 
pologist S. Fedorova, and he documents that Hawai- 
ians, Japanese, Chinese and Russians presented a major 
group of settlers. In addition, the book reports a lot of 
British Columbian and Vancouver history and puts 
Canada in the context of the overall Pacific. 

Despite the fact that whaling, sealing and eating 
dolphins is as old as the human history of the North 
Pacific, whale watching (starting as early as 1907) has 
already produced more profit than commercial whal- 
ing ever did for western North America. Greenpeace 
started in Vancouver, it “was born in the blood of 
whales”. Nevertheless, the author shows that already 
in the 19" century the pelagic seal hunt provoked the 
first great international controversy about the over- 
harvesting of the world’s marine mammals. It resulted 
in the international milestone contract (“‘fur seal treaty”’) 
of 1911 between Russia, Japan, Canada and USA. 

Topics mentioned in the seven chapters of this book 
are so manifold and detailed that only some can be 
mentioned in this review: Bute Wax, Russian scientist 
K. V. Belkemishev, occurrence of pilchards in British 
Columbia, oolichan grease, geoduck, Pollock fisheries, 
Korean squid fisheries, canneries, Earth Rotational Vel- 
ocity Index, J. Cook, G. Vancouver, V. Bering and J. J. 
Walbaum. Although the author emphasizes the prob- 
lems with old-fashioned type of science for the North 
Pacific, the book is actually based on scientific publica- 
tions. The index and the annotated scientific references 
will be highly appreciated by the scholar. This text book 
(no pictures but five maps) has no shortcomings. 


FALK HUETTMANN 


Biology and Wildlife Department, University of Alaska, Fair- 
banks, Alaska 99775-7000 USA 


Quantitative Conservation Biology: Theory and Practice of Population Viability Analysis 


Morris, W. F., and D. F. Doak. 2002. Sinauer Associates Inc., 
Sunderland Massachusetts USA. ISBN 0-87893-546-0 
paperback (41.95 US$) 


This is a great book, which should affect how we 
research and manage wildlife and its controlling fac- 
tors. The topic of a Population Viability Analysis 
(PVA) is not really new, but there are only few books 
that describe the topic well for the general public and 
managers. “PVA is the use of quantitative methods to 
predict the likely future status of a population or col- 
lection of populations of conservation concern”. 


“The promise that PVA holds as a tool for guiding 
conservation decision-making has been recognized by 
governmental science advisory boards, by professional 
organizations such as the Ecological Society of Amer- 
ica and by nongovernmental conservation organiza- 
tions such as The Nature Conservancy.” This statement 
also holds for the Habitat Conservation Plans and for 
the Recovery Plans of the U.S. Endangered Species 
Act. However, “Instead of seeing PVA as a valuable 
tool to aid their decision making, most field-oriented 
conservation biologists retain the misinterpretation 


2005 


that PVA models can only be constructed and under- 
stood by an elite priesthood of mathematical popula- 
tion ecologists”. 

Fortunately, this book is supposed to make PVAs 
easier to understand. It is based on the advanced 
matrix-based population modeling concept and uses 
count-based and demographic PVAs. The authors pres- 
ent actually a very good introduction to demographical 
population studies and even to the relatively new AIC 
concept. It explains its concepts with examples from a 
great variety of different animal and plant populations 
world-wide. The authors do a great effort to explain 
important concepts such as Vital Rates, Lambda, Bon- 
anzas and Catastrophes, Density Dependence, Ricker 
Curve, Beverton-Holt Model, Log-Population Growth 
Rate, Accounting for Errors, Environmental Stochas- 
ticity, Sensitivity Analysis and many others. As a key 
take-home message from this book I see the authors’ 
focus on confidence intervals, rather than the pure pop- 
ulation means. Such an approach embraces the uncer- 
tainty among population estimates in a much more 
transparent fashion than usually done. Many conser- 
vationists world-wide have encountered the sad but 
so often true statement made by the authors: “While 
data uncertainties are frequently used as a reason to 
rely solely on expert opinion — or on simple political 
expediency — when deciding difficult issues, we believe 
that use of more formal analyses can frequently benefit 
conservation practice. In the absence of such scientific 
analysis of conservation situations, personalities, pol- 
itics, and dollars will drive what actions are and are not 
taken, often with little or no regard to their real con- 
servation value”. 

The reader will also learn in this excellent PVA-book 
about the great importance of the extinction-time cumu- 
lative distribution function, plotted against years into 
the future. As the authors show, there are five meas- 
ures to express extinction risk: the probability of extinc- 
tion by a given time, the probability of extinction ever 


BoOoK REVIEWS 


occurring, and the mean, median and model times to 
extinction. Of these, only the first three are the most 
useful, but the last two are still the ones most often 
used. 

This book has contributing software in MATLAB 
and SAS code (also available on the website www. 
sinauer.com/PVA/), which the practitioner will bene- 
fit from. Fourteen pages of literature references and a 
well-organized index will be very helpful to the reader 
as well. 

Despite the “how to” focus of the book, I find the 
text is not that easy to understand, and it refers the 
reader too often all over the book. So from my expe- 
rience, I suspect that most managers will not really 
read it, nor fully understand all relevant (statistical) 
details; the mathematical codes alone take up an 
Appendix. The book on how to link PVAs with Geo- 
graphic Information Systems (GIS) still awaits to be 
written. 

In either case, I admire in this book that is promotes 
an overall quantitative approach to wildlife conserva- 
tion, and specifically I love the last chapters; e.g., Man- 
agement with Uncertainity, Multiple Site PVAs, Via- 
bility-Analysis for Spatially Structured Populations and 
When and When Not to Perform a PVA (a great argu- 
mentation help when doing PVAs). There just is no 
escape from numbers and reliability in this important 
conservation field. 

This important book makes it clear that well-des- 
igned demographical studies and PVAs are nowadays 
among the basics for any wildlife population to be 
studied and managed. It provides crucial tools for a 
quantitative wildlife monitoring and conservation in 
the new millienium. Now it’s once more up to the man- 
agers to read, to understand, and fully implement all 
relevant lessons learnt from this baseline publication. 


FALK HUETTMANN 


Biology and Wildlife Department, University of Alaska, Fair- 
banks, Alaska 99775-7000 USA 


Their Fathers’ Work: Casting Nets with the World’s Fishermen 


By W. McCloskey. 2000. International Marine/McGraw-Hill, 
P.O. Box 182604, Columbus, Ohio 43272 USA. 370 pages, 
$20.95 Paper. 


This book provides the reader with a superb and 
highly praised overview of global fisheries, focusing 
on Alaskan waters. In addition, it also covers first-hand 
experiences for offshore and coastal fisheries with 
vessels from Japan, Chile, Indonesia, Newfoundland 
(Grand Banks), Maine (Georges Bank), Iceland and 
Norway. The book is very pleasant to read since it 
combines fiction with facts. It is a heroic and roman- 
tic description of a likely soon-to-be-gone life of 
hard work. Nevertheless, reading how other people 
work very hard and under life-threatening conditions 
might present some sort of decadence; but so be it. 


In case the reader would not be familiar with how 
to cheat in the business of international fisheries and 
quotas (led by Spain, Taiwan, Japan and many East 
European nations) this book will definitely help. It out- 
lines in detail how fishing quotas are easily doubled, 
if not ignored by many vessel captains and fishermen 
worldwide. The explicit use of Dynamite Fishing, Liner 
Nets (an additional net with an illegally smaller mesh- 
size put inside the regular net), the “Pareja” Method 
(one huge net pulled by two boats) and many other 
tricks are shown and suggested; e.g., the same vessel 
being registered with two different names (thus, mul- 
tiplying the quota by two), stowing an additional catch 
somewhere under deck, trading the catch offshore 
(therefore enabled to start again with a “new” quota), 


316 


and mis-reporting catches. When fisheries officers ap- 
pear for control and gear inspection, nets simply get 
cut off (which makes it even worse for fish, seabirds 
and sea mammals that drown in the “ghost nets” later). 
Overall, I find that the author, an American, might have 
a tendency to blame the Spaniards and Russians too 
much here. Instead, a mention and description of the 
role that the Vladivostok-based Russian fisheries plays, 
acting worldwide, could have made the book even better. 

The thorough understatement of environmental dam- 
age done by coastal and offshore fishery must be of 
concern to any informed naturalist. The author neglects 
to address the destructive fishery method from draggers 
(“seafloor dredging”), which is, for instance, estimated 
to damage an area larger than that lost through defor- 
estation in the tropics. There is no mentioning of fish- 
eries gear polluting beaches worldwide, or “ghost nets” 
which float around in the world’s oceans for years 
(eventually, they will sink, but only the fish know 
whether they will ever rot). Sensitive by-catch topics 
such as the endangered Short-tailed Albatross (Phoe- 
bastria albatrus) caught by freezer-longliners fishing 
off Alaska are not mentioned, and certainly there is no 
reporting of the numerous sea turtles, sharks, dolphins, 
porpoises, seabirds, moon fishes and many other spe- 
cies suffering and dying for the sake of high quality 
fish. In times of environmentalism, that might be seen 
as a short coming of this book. Although the occur- 
rence of a “black catch” is somewhat mentioned, one 
has to read that shrimp fisheries has apparently almost 
no by-catch. The reader has to keep his/her breath 
when McCloskey mentions “overpopulations” of Sock- 
eye and seals; 50 000 seals are described as an “‘over- 
population” rather than victims in a potential by-catch 
problem. No wonder, the author identifies clearly from 
the “fishermen’s side”, blames Greenpeace, and does 
not place fisheries in the overall context of the envi- 
ronment; instead, he mostly focuses on economical 
and descriptive aspects of fisheries. In this regard, the 
author’s presentation of Chile’s fishery development 
lacks sensitivity to the well-proven and negative effects 
of over-commercialization. On the other side, his won- 


MISCELLANEOUS 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


derful and detailed presentation of the effects from 
the Exxon Valdez Oilspill for Alaska and its island 
communities compensate for the previous short-com- 
ings. A remarkable link is shown why the prizes of 
the Japanese Salmon market are driven by cycles of the 
Japanese Salmon runs, and thus dictate the Alaskan 
Salmon fisheries. McCloskey gets closer to the heart 
of the fisheries problem when outlining that improved 
efficiency and introduction of very light, and therefore 
allowing for longer, plastic nets has contributed to 
the current overfishing crisis. 

In the numerous and fascinating book chapters the 
author also emphazises and describes that there exists 
such a thing as severe overfishing: Snow Crab in Alas- 
ka; Cod , Flounder and Squid in Newfoundland; and 
Halibut off West America. He blames governmental 
mismanagement and elaborates nicely throughout the 
text that there is also conflict of interest among fish- 
ermen on these topics; e.g., unions, and small scale fish- 
eries vs. industrialized trawlers. In the context of gov- 
ernmental mismanagement, New Zealand’s Orange 
Roughy, a prime example of overfishing and disas- 
trous fisheries management, could have been men- 
tioned, too. The book would have gotten even better 
when topics such a Native Fishery Rights, North Sea 
Fisheries and Krill Fisheries in the Antarctic would 
have been included. The map of the Grand Banks lacks 
the French Fisheries zone around St. Pierre and Mique- 
lon; but the reader will appreciate that this book has a 
very detailed index, which allows that it can serve as 
a valid source of references, too. 

The book ends with a well-written and conclusive 
section on global fisheries and policy. The author 
quotes from one of his many interviews with experts: 
“Gathering fishery statistics is an art in probability”. 
That statement makes it clear that, currently, there can 
be no sustainable world fisheries. Due to the many top- 
ics covered, I thoroughly enjoyed reading this book 
and got literally “hooked”. 


FALK HUETTMANN 


Biology and Wildlife Department, University of Alaska, Fair- 
banks, Alaska 99775-7000 USA 


How the Earthquake Bird Got its Name and Other Tails of an Unbalanced Nature 


H. H. Shugart. 2004. Yale University Press, New Haven, 
USA. 227 pages. 


What do the following five birds, four mammals and 
one marsupial have in common: Ivory-billed Wood- 
peckers, penguins, packrats, Bachman’s Warblers, 
Leadbeater’s Possums, Red-billed Queleas, Beavers, 
Giant Moas, Gray Wolves, and European Rabbits? 
Several are extinct, a few are very numerous, some are 
common, and others are rare. They all have been cho- 
sen by Shugart who, with charm and panache, intro- 


duces the reader to a wide range of ecological con- 
cepts under the rubric of animal parables. 

Shugart, the W. W. Corcoran Professor and Director, 
Global Environmental Change Program at the Uni- 
versity of Virginia, presents nine ecological concepts: 
forest gap dynamics, niche theory, paleoecology, eco- 
logical disturbance, migration, keystone species, island 
biogeography, domestication, and invasive species. 
These ecological principles are not presented in a 
“pristine” form, but are embedded within the context 
of human transformation of the earth’s landscapes and 


2005 


how these transformations relate to animal extinc- 
tions and explosions. 

Shugart introduces each concept with an animal 
story that sets the stage for an intelligent and enter- 
taining journey through a mélange of natural and 
human history. Did you know that the word “pen- 
guin” comes from two Welsh words? Pen is the Welsh 
word for head and gwyn is Welsh for white. Penguins 
do not have white heads, but were actually named for 
the great, guano-whitened headlands on an island 
near Newfoundland (Funk Island). The birds of inter- 
est, however, were not penguins as we know them 
from the Antarctic region, but actually auks, of which 
the now extinct Great Auk was the one most familiar 
to the sailors of the day. The Great Auks were named 
penguins long before European mariners misnamed 
the similar looking but unrelated penguins of the south- 
ern and Antarctic waters. Being from Newfoundland, 
I found this little tidbit delightful. This is typical of 
the manner in which Shugart expertly weaves natural 
and human history into an attractive and colourful 
mosaic. 

The packrat and its middens introduce the reader 
to the world of paleoecology, of the archival nature 
of tree rings, ice cores, and pollen deposits that reveals 
a dynamic and ever-changing earth. The African grass- 
land Red-billed Quelqua, apparently the most com- 
mon bird on earth, initiates the reader to a marvelous 
treatment of bird migration that is sobered by an ac- 
count of the extinction of the once numerous Passenger 
Pigeon. Shugart’s account of the Wolf (Canus lupus) 
introduces us to the domesticated wolf or dog (Canus 
domesticus) and the history of animal domestication 
by our ancestors. To see how humans have radically 


NEw TITLES ;Available * Assigned 


Zoology 


Amphibians and Reptiles of the Bay Islands, and Cayos 
Cochinos, Honduras. By J. McCranie, L. Wilson and G. K6h- 
ler. 2005 Bibliomania! P.O. Box 58355, Salt Lake City, Utah 
84158. 224 pages. U.S. $29.95 Cloth. 


Amphibians and Reptiles: Status and Conservation in 
Florida. 2005. By W. Menshaka and K. Babbitt. P.O. Box 
9542, Melbourne, Florida USA. 32902-9542. 334 pages. 
U.S. $66.50. 


Bird Coloration — Volume 1. By G. Hill and McGraw. 2005. 
| Harvard University Press, 100 Maple Ridge Drive, Cumber- 
} land Rhode Island 02864-1769. 544 pages. U.S. $95. 


| * Birds of New Brunswick: An Annotated list. (Oiseaux 

du Nouveau-Brunswick: une liste commenteé.) By David 

Christie et al. 2005. New Brunswick Museum, Monograph 

| No. 10, 277 Douglas Avenue, Saint John, New Brunswick 
E2K IES. 84 pages. not illustrated, no price available. 


Book REVIEWS 


317 


altered natural landscapes without metal or modern 
devices, one has only to turn to the domesticated graz- 
ers. 

If you are looking for support of notions such as 
“the balance of nature” or “unspoiled, pristine wilder- 
ness” you will be disappointed. Shugart explicitly 
eschews such notions. His objective is to “provide an 
alternative view, to give insights into the dynamically 
changing nature of ecosystems and the implications 
of this dynamism for our stewardship of the planet” 
(page 2). According to Shugart, the only constant in 
nature is change. For him, planetary management or 
stewardship is the human vocation; a vocation that is 
defined, not by hubris, but rather by an acknowledge- 
ment of the long and continuing history of human 
alteration of planetary ecosystems, and the need for 
intentional and responsible human action. It is within 
this paradigm that we must understand the myriad of 
conservation and ethical challenges that face us. 

This work will pique the interest of all naturalists. 
Shugart’s writing is far from being pedantic or stogy. 
He writes with passion, charm and clarity about a sub- 
ject that has no doubt become a vocation. A wealth of 
original ecological research is synthesized in a delight- 
fully accessible manner that relates to our proverbial 
interest in the wax and wane of animal species. Why 
are some species abundant and others rare? Why does 
one species response positively to human influence, 
while other species meet extirpation or extinction? 
The detailed and helpful notes serve well the interest- 
ed reader who wishes to pursue further research. 


JOHN MCCARTHY 


Holy Rosary Parish, 175 Emma Street, Guelph, Ontario 
NIE 1V6 Canada 


Birds of Ontario: Habitat Requirements, Limiting Factors 
and Status (Nonpasserines, Waterfowl through Cranes). 
By. A. Sandilands. 2005. UBC Press, 2029 West Mall, Van- 
couver, British Columbia VST 1Z2. 366 pages. $95. 


Birds of Two Worlds — The Ecology and Evolution of 
Migration. Edited by Russell Greenberg and Peter P. Marra. 
The Johns Hopkins University Press, 2715 North Charles 
Street, Baltimore, Maryland 21218-4363. £15.50 Cloth. 


* Blue Grouse — their Biology and Natural History. By F. 
Zwickei and J. Bendall. 2005. NRC Research Press, M-55, 
National Research Council, Ottawa, Ontario K1A OR6. 284 
pages. $69.95 Paper. 


Crows. By Candace Savage. 2005. Greystone Books, Suite 
500, 720 Bathurst Street, Toronto Ontario M5S 2R4. 120 
pages. $27 Cloth. 


Dragonflies of Sussex. By T. A. Belden et al. 2004. NHBS 
Ltd., 2-3 Wills Road, Totnes, Devon, Great Britain TQ9 5XN. 
£7.95 Paper. 


318 


Duikers of Africa: Masters of the African Forest Floor — 
A Study of Duikers — People — Hunting and Bushmeat. 
By Vivian J Wilson. 2005. NHBS Ltd., 2-3 Wills Road, Totnes, 
Devon, Great Britain TQ9 SXN £125 Cloth. 


Florida Butterfly Caterpillars and Their Host Plants. By 
M. Minno, J. Butler and D. Hall. 2005. University Press of 
Florida, 15 NW 15" Street, Gainesville, Florida 32611. 352 
pages. US$34.95. 


* The Natural History of Bermuda. By M. Thomas. 2005 
The Bermuda Zoological Society P.O. Box FL 145 Flatts, 
Florida BX Bermuda. NPA. 


Biology of Gila Monsters and Beaded Lizards. By Daniel 
D. Beck. 2005. NHBS Ltd., 2-3 Wills Road, Totnes, Devon, 
Great Britain TQ9 5XN. £26.50 Cloth. 


Parenting for Primates. By H. Smith. 2005. Harvard Uni- 
versity Press, 100 Maple Ridge Drive, Cumberland, Rhode 
Island 02864-1769. 394 pages. US$29.95. 


Secret Weapons — Defenses of Insects, Spiders, Scorpions 
and Other Many-legged Creatures. By T. Eisner, M. Eisner 
and M. Singer. 2005. Harvard University Press, 100 Maple 
Ridge Drive, Cumberland, Rhode Island 02864-1769. 365 
pages. U.S.$29.95. 


The Smaller Majority (Frogs, insects and others). By P. 
Naskreki. 2005. Harvard University Press, 100 Maple Ridge 
Drive, Cumberland, Rhode Island 02864-1769. 364 pages. 
U.S.$35. 


+ Wheatears of the Palearctic — Ecology, Behaviour and 
Evolution of the Genus Oenanthe. By E. Panov. 2005. Pen- 
soft Publishers, Geo Milev Street 13a, 1111 Sofia, Bulgaria. 
In English, pages. U.S.$58.80. Cloth. 


+ Land Snails of British Columbia. By R. Forsythe. 2005. 
Royal BC Museum, 675 Belleville Street, Victoria, British 
Columbia V8W 9W2. 192 pages. $25.95. 


* Monitoring Bird Populations Using Mist Nets. By C. 
Ralph and E. Dunn (eds). 2005. Cooper Ornithological Soci- 
ety. 211 pages. U.S.$23 Paper 


Nests, Eggs, and Nestlings of North American Birds (Sec- 
ond Edition). By P. Baicich and C. Harrison. 2005. Princeton 
University Press, 41 William Street, Princeton, New Jersey, 
08540-5237 USA. 416 pages. U.S.$29.95 


Out of the Blue — a Journey through the World’s Oceans. 
By P. Horsman. 2005. The MIT Press, Five Cambridge Cen- 
ter, 4'" Floor, Cambridge, Massachusetts 02142-1493 USA. 
160 pages. U.S.$22.95 Cloth. 


Walker’s Carnivores of the World. By Ronald M. Nowak. 
2005. The Johns Hopkins University Press, 2715 North Charles 
Street, Baltimore, Maryland 21218-4363. £15.50 Paper. 


Botany 


Forests in Landscapes — Ecosystem Approaches to Sus- 
tainability. Edited by Jeffrey A. Sayer and Stewart Maginnis. 
2005. NHBS Ltd., 2-3 Wills Road, Totnes, Devon, Great 
Britain TQ9 5XN £29.95 Cloth. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Forest Pathology: From Genes to Landscapes. Edited by 
J. E. Lundquist and R. C. Hamelin. 2005. The American Phy- 
topathological Society/APS Press, 3340 Pilot Knob Road, 
Saint Paul, MN 55121. 175 pages. U.S.$69. Paper. 


Primates of Colombia. By R. T. Defler. 2005. Conservation 
International — Tropical Field Guide Series. NHBS Ltd., 2-3 
Wills Road, Totnes, Devon, Great Britain TQ9 5XN. 550 
pages. US$40.00. Paper. 


Trees — National Champions. By R. Conover. 2005. The MIT 
Press, Five Cambridge Center, 4° Floor, Cambridge, Mass- 
achusetts 02142-1493, USA. 144 pages. US$39.95. 


The Vascular Plant Red Data List for Great Britain — 
Species Status 7. Edited by C. M. Cheffings and L. Farrell 
softcover. 2005. NHBS Ltd., 2-3 Wills Road, Totnes, Devon, 
Great Britain TQ9 5XN. £15Paper or as a download see — 
http://www.jncc.gov.uk/page-3354. 


Environment 


+ Survival by Hunting — Prehistoric Human Predations 
and Animal Prey. By G. Frison . 2005 University of Califor- 
nia Press, 2120 Berkeley Way, Berkeley, California 94704- 
1012. 


Wild Prairie. By J. Page. 2005. Greystone Books, Suite 500, 
720 Bathurst Street, Toronto Ontario M5S 2R4. 144 pages. 
$45. Cloth. 


Aquatic Food Webs — An Ecosystem Approach. Edited by 
A. Belgrano, U. Scharler, J. Dunne and B. Ulanowicz. 2005. 
NHBS Ltd., 2-3 Wills Road, Totnes, Devon, Great Britain 
TQ9 5XN. £39.95 Paper 


Beyond Conservation — A Wildland Strategy. By Peter 
Taylor. 2005. Earthscan / James & James, 8-12 Camden High 
Street, London NW1 OJH, UK. £59.99 Cloth. 


* Manly Hardy (1832-1910): The Life and Writing of a 
Maine Fur-Buyer, Hunter, and Naturalist. By William B. 
Krohn. 2005. Maine Folklife Center, 5773 So. Stevens Hall, 
University of Maine, Orono, Maine 04469. 343 pages. US 
$24.95. Hardcover. US$19.95. Paper. 


Marine Conservation Biology — The Science of Main- 
taining the Sea’s Biodiversity. By Elliot A. Nourse and 
Larry B. Crowder. 2005. Island Press, 1718 Connecticut Ave, 
NW, Suite 300, Washington DC, 20009. £33.50 Cloth. 


Ocean and Coastal Conservation Guide 2005-2006 — The 
Blue Movement Directory. By David Helvarg. 2005. Island 
Press, 1718 Connecticut Ave, NW, Suite 300, Washington DC, 
20009. £18.50 Paper. 


Out of Gas — The End of the Age of Oil. By David Good- 
stein. 2005. NHBS Ltd., 2-3 Wills Road, Totnes, Devon, Great 
Britain TQ9 5XN. £8.99 Paper. 


Secret Nature of Devon. By Andrew Cooper. 2005. NHBS 
Ltd., 2-3 Wills Road, Totnes, Devon, Great Britain TQ9 5XN. 
£12.95 Paper. 


Spatial Analysis — A Guide for Ecologists. By Marie-Josee 
Fortin and Mark Dale. 2005. NHBS Ltd., 2-3 Wills Road, 
Totnes, Devon, Great Britain TQ9 5XN. 365 pages. £53.99 
Cloth, £26.99 Paper. 


News and Comment 


Marine Turtle Newsletter (108) 


April 2005. 36 pages: ARTICLES: Record 42 Kemp’s Ridley 
nests found in Texas in 2004 — Sea turtle strandings and 
mortality in Ecuador 1994-1999 — Observations of marine 
turtles during seismic surveys off Bahia, northwestern Brazil 
— Zakynthos sea turtle odyssey — A political ball game — 
NOTES: Hurricane effects on nesting Caretta caretta — Log- 
gerhead Turtles in the Dalyan River, Mullga Province, Turkey, 
2004 — IUCNMTSG UPDATE — MEETING REPORTS — 
BOOK REVIEWS — ANNOUNCEMENTS — NEWS & LEGAL 
BRIEFS — RECENT PUBLICATIONS. 


The Marine Turtle Newsletter is edited by Brendan J. 
Godley and Annette C. Broderick, Marine Turtle Research 
Group, Centre for Ecology and Conservation, University of 
Exeter in Cornwall, Tremough Campus, Penryn TR10 9EZ 
United Kingdom; e-mail MTN @seaturtle.org; Fax +44 1392 
263700. Subscriptions and donations towards the production 
of the MTN can be made online at <http://www.seaturtle. 
org/mtn/> or postal mail to Michael Coyne (online Editor) 
Marine Turtle Newsletter, | Southampton Place, Durham, 
North Carolina 27705 USA (e-mail: mcoyne @seaturtle.org). 


The Boreal Dip Net/L’Epuisette Boreale: Newsletter of the Canadian Amphibian and Reptile 
Conservation Network/ Reseau Canadien de Conservation des Amphibiens et des Reptiles 


9(1) April 2005 


Editor’s Note (Kerrie Serben) — Snakes on tour — CARC- 
NET and Sciensational Ssnakes!! Team up to provide public 
education (Jeff Hathaway) — South Okanagan spotlight on 
species: Tiger Salamander (Susan L. Ashpole and Peter Ord) 

-— Evaluation of the Eastern Massasauga Rattlesnake public 
education and outreach program in the Bruce Peninsula and 
eastern Georgian Bay region (Sean Liipere) — Notes from 
the field (Larry Halverson) — Herping in Costa Rica (Jonathan 
Choquette) — Advances in anuran apprehension, a technical 


note (Frederick W. Schueler) — CARCNET increases mem- 
bership fees to $29 per year (David A. Galbraith) — 9" Annual 
Living Lakes Conference (Larry Halverson) — Upcoming 
Meetings of interest — Recent articles of interest. 

Membership in CARCNET/RECCAR (contact Bruce Pauli, 
Canadian Wildlife Service, National Wildlife Research Centre, 
Carleton University, Raven Road, Ottawa, Ontario K1 A 0OH3. 
Web site: http://www.carcnet.ca/). 


Ontario Natural Heritage Information Centre Science and Information Newsletter 10(1) 


Winter 2005 


Contents of this 20 page issue: NHIC adopts BIOTICS 
software — Protecting Ontario’s biota: Your species field notes 
— Shadowdragons in the dark: Another new dragonfly for 
Ontario — NHIC staff coordinate atlassing trip to the Pen 
Islands — Riverine alvars and prairies in southern Ontario 
— NHIC produces data summaries on Niagara escarpment 
— NHIC explores electronic handheld technology — General 
status of wild species update — Species at risk element 


} occurrence records added — NHIC’s involvement in the 


Canada-Ontario agreement respecting the Great Lakes Basin 
| — Project to update species at risk data for National Parks 
— Species at risk Spotted Turtles in Muskoka and Parry 
Sound districts — Species at risk EO data polygon delineation 


‘| project — Great Lakes conservation blueprint for biodiversity 
‘| nears completion — The new species at risk (SARO) list — 


CITES Control List 2005 


Two invasive aquatic plants: Fanwort and European Frog-bit 
— NatureServe develops invasive species assessment protocol 
— NHIC participates in Canadian river heritage conference 
— Trent University internships in conservation biology — 
Prince Edward County herp data files archived in NHIC — 
NHIC a partner in northern Ontario plant database project 
— NHIC adopts NatureServe’s benchmark data content 
standards — NatureServe Canada and Canadian Wildlife 
Service join forces — sharing agreement with NatureServe 
and NatureServe Canada — Rook Reviews — NHIC staff 
information. 

Mailing address for Natural Heritage Information Centre, 
Ontario Ministry of Natural Resources, 300 Water Street, 2" 
Floor, North Tower, P.O. Box 7000, Peterborough, Ontario 
K9J 8MS5, Canada; www.mnr.gov.on.ca/MNR/nhic.cfm. 


Available on the CITES-CANADA web site www.cites.ec. gc.ca 


319 


320 THE CANADIAN FIELD-NATURALIST Vol. 119 


42™ North American Moose Conference, 12—16, 2006 Baddeck, Nova Scotia, Canada 


Nova Scotia Department of Natural Resources is hosting the North American Moose Conference and Workshop held at 
the Inverary Resort in the scenic community of Baddeck, Nova Scotia. The conference aim is to facilitate the exchange of 
scientific and experimental knowledge among Moose biologists throughout North America and beyond. 

Conference theme: 

“Management Challenges of Extremes in Population Density” 

Details at: http://gov.ns.ca/natr/mooseconference. 

Anthony (Tony) L. Nette, Chair, 2005 North American Moose Conference & Workshop, Manager, Wildlife Resources, 
Wildlife Division, 136 Exhibition Street, Kentville, Nova Scotia B4N 4E5 or call (902) 679-6140; Fax (902) 679-6176, or 
e-mail netteal @ gov.ns.ca. 


Erratum: The Canadian Field-Naturalist 119(1) 


Seasonal diets of Newfoundland Martin, Martes americana attrata JOHN W. GOSSE AND BRIAN J. HEARN 
Page 46: Addition to Literature Cited: 

Gosse, J., J. W. Cox, and S.W. Avery. 2005. Home-range characteristics and habitat use by American martins in eastern 
Newfoundland. Journal of Mammalogy 86: 1156-163. 


= 


Editor’s Report for Volume 118 (2004) 


Mailing dates for issues in volume 118 were: (1) 15 
December 2004, (2) 10 May 2005, (3), 21 August 2005, 
(4) 3 March 2006. A summary of membership and 
subscriber totals 2004 is given in Table 1. The number 
of articles and notes in volume 118 is summarized in 
Table 2 by topic; totals for Book Reviews and New 
Titles are given in Table 3, and the distribution of con- 
tent by page totals per issue in Table 4. 118(1) featured 
a lead article on the History of The Ottawa Field-Nat- 
uralist’s Club 1879-2005 prepared by Daniel F. Brun- 
ton at the request of council as part the recognition of 
125 years of the Club’s existence. I am particular in- 
debted to Frank Pope and the Publication Committee: 
Ron Bedford, Bill Cody, Fenja Brodo, Karen Hamilton, 
Elizabeth Morton, and Joyce Reddoch; for advice, assis- 
tance in choice of illustrations, and review of this paper. 
Joyce Reddoch (Trail & Landscape files) and Robert 
Lee (Macoun Field Club archives) made particularly 
major contributions to the photo selections. Tributes in 
volume 118 were to Loris Shano Russell (1904-1998), 
Thomas Henry Manning (1911-1998) and Victor Kent 
Prest (1913-2003). 

Council continued to contribute 40% of membership 
dues for publication. All of subscriptions (both individ- 
ual and institutional) also go toward publication. Coun- 
cil also has allocated 80% of the annual interest from 
the Manning Fund and other capital funds to The Cana- 
dian Field-Naturalist. Manning fund portion is specifi- 
cally to offset the publication cost of northern papers 
where author and institutional contributions were insuf- 
ficient to cover page charges. 

The journal was printed at Gilmore Printers, Ottawa, 
and thanks are due business representatives Emil Holst 
for 118(1) and initiating 118(2), and to Tom Smith for 
the remaining issues; to customer representatives Ally 
Reckzin and Chuck Graham for overseeing production 
and to Wendy Cotie for type inputting, formatting, and 
corrections for all issues. Particular tribute is due Emil 


TABLE 2. Number of articles and notes published in The Cana- 
dian Field-Naturalist Volume 118 (2004) by major field of study. 


Subject Articles Notes Total 
125 Aniversary l 0 | 

Mammals 21 12 33 

Birds 5 9 18 
Amphibians + reptiles 2 3 ) 

Fish 3 0 3, 
Invertebrates ) 2 Wl 
Plants** 12 3 15 
Multigroups a 0 2 
Tributes 3 0 3 
Totals 58 29 87 


* includes articles on evironmental change in Nova Scotia and 
introduced marine species in Haida Gwaii region in 118(1). 


Holst who was guardian and guide for Bill and I, and 
for my predecessor, Loraine Smith (CFN Editor 1972- 
1981) since the mid-1970s starting with volume 90 
through associations with MOM (formally Mail-O- 
Matic), St. Joseph, and Gilmore printers. 

Leslie Cody prepared the Index for volume 118; 
Elizabeth Morton proofed the galleys. Business Manag- 
er Bill Cody handled all liason with the printer and all 
journal business, including reprint requests and billing 
and oversaw and proofed the compilation of the Index. 
Roy John arranged for book reviews, edited them, and 
prepared the New Titles listing. 

Manuscripts (excluding book reviews, notices, and 
reports) submitted to The Canadian Field-Naturalist 
totalled 75 in 2004, down 9 from 84 in 2003. The fol- 
lowing reviewed for papers submitted in 2004 (with 
number of manuscripts reviewed in parentheses if more 
than one): Associate Editors: R. Anderson, Canadian 
Museum of Nature, Ottawa, Ontario (3); C. D. Bird, 
Erskine, Alberta (11); R. R. Campbell, St. Albert, On- 
tario; P. M. Catling, Agriculture and Agri-food Canada, 


TABLE |. The 2004 circulation of The Canadian Field-Naturalist (2003 in parenthesis). Membership totals from Annual Report 
of the Ottawa Field-Naturalists’ Club, January 2005; subscription totals compiled by W. J. Cody. Forty percent of membership 
dues and 100% of subscriptions go to publication of The Canadian Field-Naturalist. Members vote on Club affairs, subscribers 


and institutions do not. 


Canada USA Other Totals 
Memberships 
Family & individual TAP LAGI) 26 (36) 6 (7) 749 = (838) 
Subscriptions 
Individuals 165" ~(163) 63 (60) 5 (6) 253) 1229) 
Institutions 164 (157) 246 (241) 30.— (30) 440 (428) 
Totals 1046 (1115) 339° G37) 41 (43) 1422 (1495) 


~| Note: 18 countries are included under “Other” (outside Canada and United States): Australia, Belgium, Denmark, Finland, France 
(3: including | to St. Pierre & Miquelon), Germany (3), Iceland, Ireland, Japan (2), Netherlands (2), New Zealand (2), Norway 
(6), Poland, Russia, Spain (2), Sweden (2), Switzerland, and United Kingdom (10: including Scotland and Northern Ireland). 


Syl 


322 


TABLE 3. Number of reviews and new titles published in Book 
Review section of The Canadian Field-Naturalist Volume 
118 by topic. 


Reviews New Titles 
Zoology 48 69 
Botany 17 16 
Environment 12 35 
Miscellaneous 6 2 
Young Naturalists 0 4 
Totals 83 126 


Ottawa, Ontario (7); B. W. Coad, The Canadian Muse- 


um of Nature, Ottawa, Ontario (5); A. J. Erskine, Sack- 
ville, New Brunswick (21); D. F. McAlpine, New 
Brunswick Museum, Saint John, New Brunswick (7); 
D. W. Nagorsen, Mammalia Biological Consulting, 
Victoria, British Columbia (13); W. O. Pruitt, Jr., Uni- 
versity of Manitoba, Winnipeg, Manitoba (18); Others: 
W. B. Ballard, Texas Tech University, Lubbock, Texas; 
J. Bart, United States Geological Survey, Boise, Idaho; 
R. Bedford, Ottawa, Ontario; D. R. Bennett, British 
Columbia Ministry of Forests, Saanichton; J. R. Bider, 
Ecomuseum, Ste-Anne-de-Bellevue, Quebec (2); S. 
Boudrup-Nielsen, Acadia University, Wolfville, Nova 
Scotia; J. Bowman, Ontario Ministry of Natural Re- 
sources, Peterbrough, Ontario; E. L. Bousfield, Ottawa, 
Ontario; J. D. Brawn, University of Illinois, Cham- 
paignn, Illinois; I. Brodo, Canadian Museum of Nature, 
Ottawa, Ontario; M. Burt, University of New Brunswick, 
Fredericton; D. J. Buckle, Saskatoon, Saskatchewan; 
C. M. Buddle, McGill University, Macdonald Campus, 
Ste. Anne-de-bellevue, Quebec; L. Carbyn, Canadian 
Wildlife Service, Edmonton, Alberta; W. J. Cody, 
Canada Agriculture and Agri-food, Ottawa; D. Cone, 
St. Mary’s University, Halifax, Nova Scotia; M. Créte, 
Québec Société de la faune et des parcs, Québec; K. 
DeSmet, Manitoba Conservation. Winnipeg; A. W, Dia- 
mond, University of New Brunswick; L. Foote, Uni- 
versity of Alberta, Edmonton; G. Forbes. University 
of New Brunswick, Fredericton; C. M. Francis, Cana- 
dian Wildlife Service, Ottawa, Ontario; J. Gilhen, Nova 
Scotia Museum of Natural History, Halifax, Nova 
Scotia (2); P. Goossen, Canadian Wildlife Service, Ed- 
monton, Alberta; P. T. Gregory, University of Victoria, 
British Columbia; E. Haber, National Botanical Ser- 
vices, Ottawa, Ontario; P. Hamilton, Canadian Museum 
of Nature, Ottawa, Ontario; M. B. Hickey, St. Lawrence 
River Institute of Environmental Studies, Cornwall, 
Ontario; C. S. Houston, Saskatoon, Saskatchewan; R. 
James, Sutherland, Ontario (2); J. Kamler, Polish Acad- 
emy of Sciences, Mammal Research Institute, Bialo- 
wieza; R. W. Knapton, Edmonton, Alberta; J. M. Knet- 
ter, University of Wisconsin-Madison; D. Langor, 
Edmonton, Alberta; P. F. Larsen, Bigelow Laboratory 
for Ocean, West Boothbay Harbor, Maine; S. Lariviere, 
Portage La Prairie, Maanitoba; J. Leafloor, Canadian 
Wildlife Service, Winnipeg, Manitoba; L. E. Licht, 
York University, North York, Ontario; R. MacCulloch, 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE 4. Number of pages per section published in The Cana- 
dian Field-Naturalist Volume 118 (2004) by issue. 


(1) (2) (3) (4) Total 
Articles 118... 05. #133 107 465 
Notes 19 i7 L7 16 69 
Tributes 0 0 14 12 26 
Book Reviews’ ii 13 19 14 63 
CFN/OFNC Reports “* 0 2 8 3 13 
News and Comment 3 2 2 3 10 
Index 0 0 0 28 28 
Advice to Contributors | | 1 | 4 
Totals 158 140 196 184 678 


“Total pages for book review section include both reviews and 


new titles listings. 
“Includes CFN Editors’ report in 118(2), OFNC Annual Busi- 
ness Meeting 118(3) and OFNC Awards 118(4). 


Royal Ontario Museum, Toronto, Ontario; G. L. Mackie, 
University of Guelph, Ontario; C. Machtans, Canadian 
Wildlife Service, Yellowknife, Northwest Territories; 
A. Martel, Canadian Museum of Nature, Ottawa, On- 
tario; D. Mazerolle, University of Saskatchewan, Sas- 
katoon; L. D. Mech, U.S. Geological Survey, The Rap- 
tor Center, University of Minnesota, St. Paul (4); J. 
Metcalfe-Smith, National Water Research Institute. 
Burlington, Ontario; E. Morton, Masham, Quebec; J. 
S. Nelson, University of Alberta, Edmonton, Alberta; — 
J.-P. Ouellet, Universite du Quebec a Rimouski, Que- 
bec; G. Parker, Canadian Wildlife Service, Sackville, 
New Brunswick; K. Poole, Aurora Wildlife Research, 
Nelson, British Columbia; E. F. Pope, Ottawa; R. 
Poulin, University of Alberta, Edmonton, Alberta; M. 
Raine, Calgary, Alberta; R. Reeves, Okapi Wildlife | 
Associates, Hudson, Quebec (2); J. Reddoch, Ottawa, | 
Ontario; T. E. Reimchen, University of British Colum- — 
bia, Victoria: J.-F. Robitaille, Laurentian University, | 
Sudbury, Ontario; F. W. Schueler, Oxford Station, On- | 
tario; S. G. Sealy, University of Manitoba, Winnipeg, | 
Manitoba; N. Simon, Newfoundland and Labrador | 
Natural Resources, Goose Bay; B. Slough, Whitehorse | 
Yukon Territory; S. Smith, Univerisity of Toronto, | 
Ontario; K. W. Stewart, Winnipeg, Manitoba; J. The- | 
berge, Oliver, British Columbia; I. Thompson, Cana- | 
dian Forest Service, Sault Ste. Marie, Ontario; A. C. 
Thoresen, McMinnville, Oregon; S. P. Vander Kloet, | 
Acadia University, Wolfville, Nova Scotia. | 

I am also indebted to the President of the Ottawa | 
Field-Naturalists’ Club Gary McNulty and the Club | 
Council for continuing support of the journal; Chair- | 
man Ron Bedford and the Publications Committee of | 
the OFNC for editorial encouragement and support, | 
to the Canadian Museum of Nature for access to its | 
library and the facilities at the Natural Heritage 
Building, 1740 Pink Road, Gatineau [Aylmer sector], | 
Quebec, and to Joyce for everything else. 


FRANCIS R. COOK | 
Editor © 


TABLE OF CONTENTS (concluded) Volume 119 Number 2 


Notes 
Apparent predation by Gray Jays, Perisoreus canadensis, on Long-toed Salamanders, 
| Ambystoma macrodactylum, in the Oregon Cascade Range 
MICHAEL P. MURRAY, CHRISTOPHER A. PEARL, and R. BRUCE BURY 


Territorial behavior in Belted Kingfishers, Ceryle alcyon, during fall migration 
. MICHAEL J. HAMAS 


First record of age 0+ Atlantic Whitefish, Coregonus huntsmani, from the wild 
D. J. HASSELMAN, PHILLIP LONGUE, R. G. BRADFORD 


assage through a small drainage culvert by Mule Deer, Odocoileus hemionus, and other 
large mammals ALISSA KRAWCHUK, KARL W. LARSEN, RICHARD D. WEIR, and HELEN DAVIS 


300k Reviews 

ZOOLOGY: Birds of Azerbaijan — Birds of New Brunswick: An Annotated List — Fishes of the Great 
| Lakes Region Revised Edition — Seabirds and Atlantic Canada’s Ship-Source Oil Pollution — 
| Spiders of Australia: An Introduction to their Classification, Biology and Distribution — Annotated 
} Bibliography of Quaternary Vertebrates of Northern North America with Radiocarbon Dates — White 
} as a Ghost: Winter Ticks and Moose — Lewis and Clark on the Great Plains: A Natural History — 
| Waterfowl of Eastern North America — Wolves: Behavior, Ecology, and Conservation 


BOTANY: Rendezvous with the Wild: The Boreal Forest 


TNVIRONMENT: The Earth’s Blanket: Traditional Teachings for Sustainable Living —- Greenpeace: How a 
Group of Journalists, Ecologists and Visionaries Changed the World — The Last Great Sea: A Voyage 
} through the Human and Natural History of the North Pacific Ocean — Quantitative Conservation 
Biology: Theory and Practice of Population Viability Analysis — Their Father’s Work: Casting Nets 
| with the World’s Fishermen 


MiscELLANEOUS: How the Earthqake Bird Got its Name and Other Tales of an Unbalanced Nature 


Jew TITLES 


‘Jews and Comment 

Varine Turtle Newsletter (108) — The Boreal Dip Net/ L’Epuisette Boreale 9(1) April 2005 — Ontario 
| Natural Heritage Information Centre Science and Information Newsletter 10(1) Winter 2005 — 
.| CITES Control List 2005 — 42nd North American Moose Conference, 12-16 June 2006 — Erratum 
‘| The Canadian Field-Naturalist 119(1) 


ditor’s Report for Volume 118 (2004) 


z [ailing date of the previous issue 119(1): 6 June 2006 


2005 


291 


pap Bs 


294 


296 


298 
308 


309 
316 
S17 


319 


321 


THE CANADIAN FIELD-NATURALIST Volume 119 Number 2 


Articles 


The influence of thermal protection on winter den selection by Porcupines, 
Erethizon dorsatum, in second-growth conifer forests Topp N. ZIMMERLING 


Black Bear, Ursus americanus, ecology on the northeast coast of Labrador 
KEITH CHAULK, SOREN BONDRUP-NIELSEN, and FRED HARRINGTON 


Long-tailed Weasel, Mustela frenata, movements and diggings in Alfalfa 
fields inhabited by Northern Pocket Gophers, Thomomys talpoides GILBERT PROULX 


Long-tailed Duck, Clangula hyemalis, eider, Somateria spp., and scoter, 
Melanitta spp., distributions in central Alaska Beaufort Sea Lagoons, 1999-2002 
LYNN E. NOEL, STEPHEN R. JOHNSON, and GILLIAN M. O’ DOHERTY 


Winter habitat use by Moose, Alces alces, in central interior British Columbia 
GILBERT PROULX and RHONDA M. KARIZ 


Facts from faeces: prey remains in Wolf, Canis lupus, faeces revise occurrence 
records for mammals of British Columbia’s coastal archipelago 
MICHAEL H. H. PRICE, CHRIS T. DARIMONT, NEVILLE N. WINCHESTER, and PAUL C. PAQUET 


Attempted predation of a child by a Gray Wolf, Canis lupus, near Icy Bay, Alaska 
MArK E. McNay and PHILIP W. MOONEY 


Post-reproductive Pacific salmon, Oncorhynchus spp., aS a major nutrient source 
for large aggregations of gulls, Larus spp. K. S. CHRISTIE and T. E. REIMCHEN 


Microhabitat characteristics of Lapland Longspur, Calcarius lapponicus, 
nests at Cape Churchill, Manitoba CLINT W. BOAL and DAvID E. ANDERSEN 


Novel surface feeding tactics of Minke Whales, Balaenoptera acutorostrata, 
in the Saguenay-St. Lawrence National Marine Park 
KATIE J. KUKER, JORDAN A. THOMSON, and URSULA TSCHERTER 


First record of the Plains Minnow, Hybognathus placitus, in Canada 
R. M. SYLVESTER, S. E. FREELING, and C. R. BERRY. JR. 


Identification and stutus of Black Pine, Pinus nigra, and Mugo Pine, 
Pinus mugo, in Ontario PAUL M. CATLING 


A potential for the use of dragonfly (Odonata) diversity as a bioindicator 
of efficiency of sewage lagoons PAUL M. CATLING 


Invasive Scots Pine, Pinus sylvestris, replacing Corema, Corema conradii, 
heathland in the Annapolis Valley, Nova Scotia PAUL M. CATLING and SUSAN CARBYN 


Insect visitation to wildflowers in the endangered Garry Oak, Quercus 
garryana, Ecosystem of British Columbia A. L. PARACHNOWITSCH and E. ELLE 


An annotated checklist of the spiders of Newfoundland 
J. R. PICKAVANCE and C. D. DONDALE 


Additions and range extensions to the vascular plant flora of the continental Northwest 
Territories and Nunavut, Canada III WILLIAM J. Copy and KENNETH L. READING 


(continued on inside back cov 


ISSN 0008-3550 


200: 


(LAN 
ISa 


The CANADIAN 
FIELD-NATURALIST 


Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada 


Volume 119, Number 3 July-September 2005 


The Ottawa Field-Naturalists’ Club 


FOUNDED IN 1879 


Patrons 
Her Excellency The Right Honourable Michaélle Jean 
Governor General of Canada 


The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to 
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields 
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ- 
ments of high quality for living things. 


Honorary Members 


Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope 
Charley D. Bird R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr. 
Donald M. Britton Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch 
Irwin M. Brodo John M. Gillett Theodore Mosquin Dan Strickland 
William J. Cody C. Stuart Houston Eugene G. Munroe John B. Theberge 
Francis R. Cook George F. Ledingham Robert W. Nero Sheila Thomson 
Ellaine Dickson 

2005 Council 
President: Mike Murphy Ronald E. Bedford —_ Diane Kitching Louise Schwartz 
Vice-President: Gillian Marston Fenja Brodo Karen McLachalan Hamilton David Smythe 
Recording Secretary: Susan Laurie-Bourque William J. Cody David Hobden Henry Steger 
Treasurer: Frank Pope Kathy Conlan Cendrine Huemer Chris Traynor 
Past President: Gary McNulty Francis R. Cook Diane Lepage Eleanor Zurbrigg 


Stanley Rosenbaum 


To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, 
Westgate P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc @achilles.net. 
For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca 


The Canadian Field-Naturalist 


The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in 
this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. 

PAP Registration Number 9477. Canada We acknowledge the financial support of the Government of Canada through the 
Publication Assistance Program (PAP) toward our mailing costs. 


All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal 
mail. Book-review correspondence should be sent by e-mail or postal mail to the Book-review Editor. 


Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: cfn@ofne.ca 
Copy Editor: Elizabeth Morton 

Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2; (613) 759-1374 
Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5, e-mail: roy.john@pwgsc.gc.ca 


Associate Editors: Robert R. Anderson Paul M. Catling David Nagorsen 
Charles D. Bird Brian W. Coad Donald F. McAlpine 
Robert R. Campbell Anthony J. Erskine William O. Pruitt, Jr. 


Chairman, Publications Committee: Ronald E. Bedford 


Subscriptions and Membership 

Subscription rates for individuals are $28 per calendar year. Libraries and other institutions may subscribe at the rate of $45 per 
year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $28 (individual) $30 (family) $50 (sustaining) and 
$500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) 
must add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and 
Local Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries 
at $28 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be 
mailed to: The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publications 
Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: April-June 2005 (November 2006). 


Cover: The small remnant peatland on the shore of Black Lake, Gatineau Park, Quebec, twenty-six years after being flooded by 
Beavers, Castor canadensis. (Photograph by Joyce Reddoch, taken on 3 July 2006.) See the paper on the consequences 
of flooding on a small shore fen by J. M. Reddoch and A. H. Reddoch, pages 385-394. 


LIBRARY 
JAN ~ 9 2007 


The Canadian Field-Natura}WinRVvarp 
a andes Snes a eae eo 


Volume 119, Number 3 July-September 2005 


An Analysis of the Parasites of a Mid-winter Population of the 
Snowshoe Hare, Lepus americanus, on Insular Newfoundland During 
a Cyclical Peak 


K. E. BENNETT!, E. M. Bacas?, J. R. FINNEY-CRAWLEY, and M. MCGRATH? 


Department of Biology, Memorial University of Newfoundland, St. John’s, Newfoundland and Labrador AIB 3X9 Canada 

'PO. Box 43, St. Paul’s Inlet, Newfoundland AOK 4Y0 Canada 

* Corresponding author 

3 Senior Small Game and Furbearer Biologist, Wildlife Division, Department of Environment and Conservation, Govern- 
ment of Newfoundland and Labrador, P.O. Box 8700, St. John’s, Newfoundland and Labrador A1B 4J6 Canada 


Bennett, K. E., E. M. Baggs, J. R. Finney-Crawley, and M. McGrath. 2005. An analysis of the parasites of a mid-winter population 
of the Snowshoe Hare, Lepus americanus, on insular Newfoundland during a cyclical peak. Canadian Field-Naturalist 
1 19(3)? 323-329. 


A mid-winter sample of 78 Snowshoe Hares (Lepus americanus) was collected during their cyclical peak in population from 
three eco-regions (Western Newfoundland, North Shore and Avalon Forest) on insular Newfoundland and was examined for 
the presence of enteric parasites. The length of the hares was significantly shorter in the Avalon Forest Region (n = 27) than 
those of the Western Newfoundland Region (n = 25) and North Shore Region (n = 26) samples (Ps 0.001 and P < 0.003 
respectively); however, no significant differences occurred for other morphological measurements. Four species of para- 
sites, two cestodes (Mosgovoyia pectinata and Taenia pisiformis) and two nematodes (Obeliscoides cuniculi and Rauschia 
triangularis), were recovered. Taenia pisiformis was recovered from the North Shore eco-region only. No other significant 
differences with respect to their prevalence, intensity, mean intensity, relative density and dispersion between eco-regions 
were found. Within eco-regions, only R. triangularis showed a significantly higher value (P< 0.027) for males and the 
prevalence of this species was lower than that previously reported. The occurrence of O. cuniculi was significantly different 
between the higher weight classes of hares and the prevalence of this species was higher than that previously reported. No 
trends for multiple infections were noted. The expansion of a new animal species, the Coyote, Canis latrans, to Newfound- 
land appeared to have had no effect on the diversity of parasites found in the hare. 


Key Words: Lepus americanus, Snowshoe Hare, parasite diversity, cestodes, Mosgovoyia pectinata, Taenia pisiformis, nema- 
todes, Obeliscoides cuniculi, Rauschia triangularis, eco-regions, exotics, insular Newfoundland. 


Following the Wisconsinian glaciation, the mam- 
malian fauna of insular Newfoundland was impover- 
ished (Dodds 1957 Scruton et al. 1995) and the island 
had a high predator to prey species ratio. Much of the 
present biodiversity is a result of accidental or deliber- 
ate introductions and colonization. The original small 
mammal (prey) populations consisted of the Arctic 
Hare (Lepus arcticus) and the Meadow Vole (Microtus 
pennsylvanicus) while the mammalian predator list 
consisted of the Lynx (Lynx canadensis), Black Bear 
(Ursus americanus), Newfoundland Wolf (Canis lupus 
beothucus, now extinct), Red Fox (Vulpes vulpes), and 
a variety of mustelids; i.e., the Pine Marten (Martes 
americana), Short-tailed Weasel (Mustela erminea), 
and River Otter (Lutra canadensis). There have also 
been temporary invasions of the Arctic Fox (Alopex 
lagopus) (Banfield 1987). 

Over the last 150 years there have been a number of 
deliberate introductions that have resulted in successful 


establishment of populations in insular Newfoundland. 
These included the Snowshoe Hare (Lepus ameri- 
canus) in 1864 and Moose (Alces alces) in 1904 (Ban- 
field 1987), which were introduced as a means of 
increasing the availability of fresh meat to the resi- 
dent human population. Small mammal introductions 
of the Red Squirrel (Jamiasciurus hudsonicus) in 1963 
(Payne 1976) and Eastern Chipmunk (7amias striatus) 
in 1962 (Northcott et al. 1973) were made to enhance 
the small mammal prey base of the island. The Deer 
Mouse (Peromyscus maniculatus), introduced in 1968 
(Gould and Pruitt 1969), is thought to have been 
accidental, imported with cargo shipped through the 
Maritime provinces of Canada to Newfoundland. The 
Red-backed Vole (Clethrionomys gapperi) (first re- 
corded about 1997 but precise year of introduction 
unknown) remains an enigma. It has been suggested 
that it established populations in the Stephenville area 
and elsewhere on the island and may have originated 


323 


324 


from transportation of logs to the liner board mill in 
Stephenville from Labrador from as far back as the 
late 1960s. Shipments of these logs were sporadic over 
that period. The Masked Shrew (Sorex cinereus) was 
introduced in 1958 in an attempt to control outbreaks 
of the Hemlock Looper (Lambdina fiscellaria) (Warren 
1970). The carnivore list increased in May 1987 when 
the Eastern Coyote (Canis latrans) became well-estab- 
lished after crossing the Gulf of St. Lawrence on win- 
ter ice floes from the Maritimes (Parker 1995). 
Introductions of exotic animals have the inherent risk 
of bringing accompanying parasites and/or pathogens 
which may subsequently impact on resident species. 
Dodds and Mackiewicz (1961) examined 630 Snow- 
shoe Hares in an attempt to establish the parasite fauna 
of these herbivores in Newfoundland and reported the 
following enteric parasites: trematode: Dicrocoelium 
dentriticum; cestodes: Mosgovoyia pectinata, Hydatig- 
era taeniaeformis, Multiceps sp. and Taenia pisiformis; 
nematodes: Trichostrongylus axei and Obeliscoides 
cuniculi. Smith and Threlfall (1972) examined three 
Snowshoe Hares and reported the presence of the ces- 
todes Multiceps serialis and Taenia pisiformes in the 
coelomic cavities, the nematode Rauschia triangularis 
in the digestive tract, and an unidentified nematode 
from the lungs. While there is no information on the 
parasite burdens of the Snowshoe Hares used in the 
1864 introductions, the work of Dodds and Mackie- 
wicz (1961) and Smith and Threlfall (1972) form a 
basis from which a comparison of the Snowshoe Hare 
parasites may be made in light of subsequent mam- 
malian introductions. This paper examines the mid- 


g 


—» 7 


300 km 3 


FIGURE |. Map of insular Newfoundland, Canada. Collection 
Areas 1, Western Newfoundland, (57°45'W, 49°6'N), 
2, North Shore (53°53'W, 48°41'N), 3, Avalon Forest 
(53°12'W, 47°30'N). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


winter enteric parasite standing of Lepus americanus 
from insular Newfoundland collected during the hare 
cyclical peak (Reynolds et al. 2004*). 


Materials and Methods 

A total of 78 adult Snowshoe Hares was collected by 
shooting and snaring by hunters and trappers from three 
eco-regions (Figure |): Western Newfoundland (Eco- 
region |), Sub-region la — Old Man’s Pond (n = 25); 
North Shore (Eco-region 3) (n = 26) and Avalon Forest 
(Eco-region 5) (n = 27) of insular Newfoundland 
(Meades and Moores 1994) and were submitted to the 
Inland Fish and Wildlife Division of the Government 
of Newfoundland and Labrador. The three eco-regions 
have similar populations of plants and animals but dif- 
fer in physical attributes and land usage. Collections 
were carried out during January through February 2000 
at the cyclical peak for Snowshoe Hares (Reynolds et 
al. 2004*) in Newfoundland. Morphological criteria 
were obtained prior to necropsy. Hares were examined 
for helminth parasites using conventional parasitologi- 
cal techniques. Descriptive statistical analysis of hare 
morphology, parasite parameters or both was accom- 
plished using the Microsoft Excel 98 statistical pack- 
age. A comparison of body length-weight relationship 
was carried out using the general linear model and a 
comparison of the slopes was made to determine 
whether the growth curves differed by eco-region. 
Comparisons of hare population data were carried out 
using standard paired t-tests and one-way ANOVAs. 
Comparisons of hare data based on sex were carried 
out using Mann-Whitney and Kruskall-Wallis non- 
parametric tests. Parasite parameters (1.e., prevalence, 
abundance etc.) were determined using the methods of 
Margolis et al. (1982). 

Data for all parasites except Rauschia triangularis 
were based on counts, but, due to the large numbers 
observed, values for R. triangularis were ranked. In all 
cases this parasite exceeded 50 individuals and a scale 
of 0 through 3 was used to rank estimated population 
numbers (i.e., 0 = 0 parasites, | = 1 — 50 individuals, 
2 = 50 — 100 parasites and 3 = more than100 individ- 
uals). 


Results 

The Snowshoe Hares had an overall sex ratio of 1.1: 
1.0 (males: females). Based on weight, there were no 
significant differences between animals from the three 
eco-regions (Table 1). There were no differences be- 
tween the body length of North Shore and Western New- 
foundland hares; however, those of the Avalon Forest 
were significantly shorter (PS 0.001 and Ps 0.003 
from North Shore and Western Newfoundland). There 
was a trend for the hares to be longer on an east to 
west basis by eco-region, although no significant dif- 
ferences were noted for any other physical character- 
istic measured. No significant differences were found 
in body length-weight relationships (P = 0.05) and 
the subpopulations were considered similar. 


2004 BENNETT, BAGGS, FINNEY-CRAWLEY, AND MCGRATH: PARASITES OF SNOWSHOE HARE 


TABLE |. Summary of morphometric data (means) on Lepus americanus from three Eco-regions on insular Newfoundland: 


Western Newfoundland, North Shore and Avalon Forest. 


Parameter 


Ecoregion 


Western Newfoundland (57°45'W, 49°6'N) 


Male (n = 12) Female (n = 12) Total (n = 24) 
Weight (kg) (+ SD) 1.35 + -0.134 1.46 + -0.125 1.405 + -0.138 
Length (mm) 442.8 + -21.70 453.3: -11.02 448.05 + -17.66 
LHFEL (mm s.u.!) 127.7 = -4.73 125.9 + -4.50 126.8 + -4.62 
Pinnae (mm) Si 2a =o > 85.8 + -4.43 86.5 + -4.24 
Slot (mm) 1032-253 70.4 + -4.06 1035653 34 

North Shore (53°53'W, 48°41'N) 

Male (n=13) Female (n=12) Total (n=25) 
Weight (kg) (+ SD) 1.34 + -0.174 1.44 + -0.133 1.39 + -0.161 
Length (mm) 440.8 + -15.77 450.6 + -14.95 445.70 + -15.86 
LHFEL (mms.u.!) 122.4 + -5.88 122 =16:06 124.80 + -11.72 
Pinnae (mm) 87.5 + -5.06 89.9 + -3.29 88.7 + -4.38 
Slot (mm) Tia +541 13:4 -9.12 12.55 + -4.45 

Avalon Forest (53°12'W, 47°30'N) 

Male (n=14) Female (n=13) Total (n=27) 
Weight (kg) (+ SD) 1.35 + -0.114 1.41 + -0.145 1.38 + -0.130 
Length (mm) 426.7 + -25.92 429.9 + -21.55 4278.30'+:-23.52 
LHFEL (mm s.u.!) 124.8 + -6.86 125.5 + -9.79 125.15 + -8.24 
Pinnae (mm) 86.8 + -6.22 88.1 + -6.28 87.45 + -6.16 
Slot (mm) 12.0% =2.33 68.5 + -4.32 70.25 + -3.65 


'L H FL —Left Hind Foot Length; s.u. — sans ungus 


TABLE 2. Occurrence of the parasites of Lepus americanus from three Eco-regions on insular Newfoundland: Western New- 


foundland, North Shore and Avalon Forest. + = present; — = not present 

Ecoregion 
Parasite Western Newfoundland North Shore Avalon Forest 
Mosgovoyia pectinata + + & 
Taenia pisiformis = re = 
Obeliscoides cuniculi + + + 
Rauschia triangularis + + + 


Sixty percent of the hares contained at least one para- 
site species, 25.6 percent contained two parasite species 
and 3.8 percent contained three parasite species. No 
hares contained four parasite species and 10.3 percent 
of the hares contained no enteric parasites. 

Four enteric parasite species were recovered. Two 
cestodes: adult Mosgovoyia pectinata and the cysticer- 
cus Taenia pisiformis and two nematodes: Obeliscoides 
cuniculi and Rauschia triangularis (Table 2). Taenia 
pisiformis occurred in one female hare in the North 
Shore Eco-region whereas M. pectinata and the two 
nematodes O. cuniculi and R. triangularis occurred in 
all three eco-regions. The parasite parameters for Lepus 
americanus from the three eco-regions are shown in 


Table 3. There were no significant differences (P = 0.05) 
for M. pectinata by sex within or between eco-regions. 
This was also the case for O. cuniculi. Raushia triangu- 
laris showed significantly higher (PS 0.027) values for 
males in the North Shore by sex, however, no differ- 
ences existed on an eco-regions basis (P= 0.05). 

An evaluation of parasite prevalence etc., for all the 
hares based on weight categories (Table 4) showed no 
significant difference (P = 0.05) between hare weight 
classes in infections of Mosgovoyia pectinata and 
Rauschia triangularis. Infections with Obeliscoides 
cuniculi showed significant differences between weight 
class 1.32 to 1.48 kg and 1.64 to 1.80 kg (Ps 0.04) and 
between weight class 1.48 to 1.64 kg and 1.64 to 1.80 kg 


326 THE CANADIAN FIELD-NATURALIST Vol. 119 


TABLE 3. Parasite parameters for Lepus americanus from three Eco-regions on insular Newfoundland: Western Newfoundland, 
North Shore and Avalon Forest. 


Species Prevalence 
W. NF (n=25) N. Shore (n=26) Av. For. (n=27) Total (n=78) 
M. pectinata 0.040 0.192 0.037 0.090 
T. pisiformis 0.000 0.038 0.000 0.013 
O. cuniculi 0.350 0.846 0.741 0.654 
R. triangularis 0.440 0.423 0.593 0.48 
Intensity 
M. pectinata Me 1-6 2 1-6 
T. pisiformis 0 l 0 l 
O. cuniculi 1 - 189 1-237 1 - 78 ] - 237 
R. triangularis 3 a af af 
Mean Intensity 
M. pectinata 1.000 2.500 2.000 2.170 
T. pisiformis 0.000 NA 0.000 NA 
O. cuniculi 29.78 29.00 32.40 28.82 
R. triangularis NA NA NA NA 
Relative Density 
M. pectinata 0.040 0.480 0.074 0.195 
T. pisiformis 0.000 NA 0.000 NA 
O. cuniculi 10.72 24.53 24.00 18.84 
R. triangularis NA NA NA NA 
Dispersion 
M. pectinata 1.000 3.970 2.000 
O. cuniculi 134.99 107.38 25.84 


Note: In all cases counts for Rauschia triangularis exceeded 50 individuals. 


W. NF — Western Newfoundland 

N. Shore — North Shore 

Av. For. — Avalon Forest 

3* sample has greater than 100 parasites 
NA — not applicable 


(P< 0.03), the majority of the parasites occurring in the 
smaller hares. The largest of the hares (n = 3) had the 
least number of parasites. 

In the smaller hares, there is a tendency for Rauschia 
triangularis to occur at a higher prevalence than Obelis- 
coides cuniculi whereas the reverse 1s seen for the heav- 
ier two weight classes (Table 4). 


Discussion 

The most extensive study of the parasites of Lepus 
americanus on the island of Newfoundland (Dodds and 
Mackiewicz 1961) involved collections of hares from 
1954 through 1959. Of a total of 630 hares they exam- 
ined, 483 were collected from the Humber East, White 
Bay South, St. Barbe and Grand Falls districts. The 
remaining 147 were collected from Emberly Island in Pla- 
centia Bay, Newfoundland, a population that was estab- 
lished in 1954 using hares from the main island. Smith 


and Threlfall (1972) reviewed the parasites of only three 
adult hares originating from the Avalon Peninsula of 
Newfoundland. 

Our study differs from Dodds and Mackiewicz (1961). 
They reported on hares from different seasons and years 
and for juveniles and adults. Our study used hares cap- 
tured during mid-winter (January/February) and exam- 
ined adults only. While containing host specimens from 
the same area as that evaluated by Dodds and Mack- 
iewicz (1961), it extends the geographic coverage of the 
island by sampling the eastern portion of the province. 

The four parasite species recovered in this study were 
the same as previous studies in the area. Dodds and 
Mackiewicz (1961) found that the prevalence of the 
cestode Mosgovoyia pectinata varied over four years 
starting with 7% prevalence in 1956, rising to a high of 
36% in 1958, and dropped to 4% in 1959. The high 
values found in 1958 occurred during the cyclical 


2004 BENNETT, BAGGS, FINNEY-CRAWLEY, AND MCGRATH: PARASITES OF SNOWSHOE HARE 


327 


TABLE 4. Parasite parameters for Lepus americanus from insular Newfoundland placed in five weight classes. 


Weight Class (kg) 


1.00-1.15 1.16-1.32 
(n= 3) (n= 18) 

Species 
M. pectinata 0.04 0.05 
T. pisiformis 0.00 0.00 
O. cuniculi 0.60 0.61 
R. triangularis 0.80 0.81 
M. pectinata 2 6 
T. pisiformis 0 0 
O. cuniculi 17-237 1-58 
R. triangularis 30 Cu 
M. pectinata 2 6 
T. pisiformis 0) 0 
O. cuniculi 105.67 20.82 
R. triangularis NA NA 
M. pectinata 0.80 0.33 
T. pisiformis 0.00 0.00 
O. cuniculi 63.4 F272 
R. triangularis NA NA 
M. pectinata | a 6.00 
T. pisiformis NA NA 
O. cuniculi 159.02 22.28 
R. triangularis NA NA 


3*— sample contains more than 100 parasites 


1.33-1.48 1.49-1.64 1.65-1.80 
(n =33) (n= 17) (n = 3) 
Prevalence 
0.03 0.06 0.00 
0.63 0.76 0.00 
0.39 0.53 0.06 
0.12 0.00 0.12 
Intensity 
I | 0) 
0 a 0 
1-189 1-78 8 
Mean Intensity 
l l 0 
0 NA 0 
28.52 22.46 0.47 
NA NA NA 
Relative Density 
0.03 0.12 0.00 
0.00 NA 0.00 
18.15 17.18 0.47 
NA NA NA 
Dispersions 
1.00 0.94 NA 
NA NA NA 
90.43 29.08 8.00 
NA NA NA 


Note — The sample size (n = 76) for weights is less than the total (n = 78) used for parasites due to heads of the hares being 


missing in two samples. 


decline in hare populations and were from juveniles 
only; adults did not contain this parasite. Dodds and 
Mackiewicz (1961) reported that their juveniles often 
presented distended small intestines which appeared 
to be occluded, although the hares otherwise appeared 
to be in good condition. In contrast, the 8% prevalence 
for M. pectinata in our study was for adults taken in 
mid-winter only and no significant differences in preva- 
lence were found between weight classes (Table 4) of 
the hares. There were no indications of occlusions or 
organ damage and the hares appeared to be in excel- 
lent condition. 

The cysticerci of Taenia pisiformis were recovered 
from only one female hare in the North Shore Ecore- 
gion, however, four out of six specimens of hares snared 
in the Avalon Forest Eco-region in 2001 contained mas- 
sive infections of 7. pisiformis (E.M.B.). Dodds and 
Mackiewicz (1961) report relatively heavy prevalences 
for T. pisiformis and Hydatigera taeniaeformis and 
lump the data to show prevalences which range from a 
high of 61% in 1956 to a low of 17% in 1958. We found 


no indication of H. taeniaeformis in this study. The rar- 
ity of T. pisiformis could be related to the scarcity of 
canids in the respective eco-regions as the definitive 
hosts for this parasite are known to be canids (i.e., Red 
Fox, Coyote and Domestic Dog) (Whitney 2000*). 
However, the eco-regions utilized in this study are areas 
where active trapping of furbearers takes place and dur- 
ing peak years of the hare cycle, there may be a dilution 
effect such that the definitive host populations would 
be low relative to the intermediate host population. Win- 
ter conditions would remove the hares from normally 
exposed vegetation on which tapeworm eggs would be 
deposited and coupled with higher mortality of such 
eggs, a loss in parasites could be expected. These low 
numbers could also be a result of conferred immunity 
from earlier infections. Heath (1973) showed immunity 
to T. pisiformis eggs in New Zealand White Rabbits fol- 
lowing injections with antigens from various stages of 
parasite development (i.e., oncospheres, larvae, etc.). 
Rickard and Coman (1977) further demonstrated that 
immunity and enhanced development of T: pisiformis 


328 


in New Zealand White Rabbits depending on which 
species of Taenia was utilized as the antigenic source. 

Obeliscoides cuniculi was more prevalent (65%) than 
any of the values reported by Dodds and Mackiewicz 
(1961). They reported a prevalence of 44% in 1956 ris- 
ing to a high of 63% in 1958, subsequently dropping to 
54% in 1959. Most of their hares were obtained during 
the summer of a cyclical peak in population. Erickson 
(1944) also reports the highest frequency for this para- 
site at the beginning of a decline in hare numbers. The 
values for O. cuniculi in this study mirror the findings 
of Erickson (1944). Keith et al. (1985) report signifi- 
cantly higher values (P = 0.07 and P =-0.02) for this par- 
asite during mid-winter and during repetitive cyclical 
peaks in 1962 and 1971 respectively. Keith et al. (1985) 
attribute the higher numbers of the winter populations 
of O. cuniculi, to the rise in the pituitary gonadotro- 
phins and testicular weights prior to the March breed- 
ing season. The cyclical increases in O. cuniculi, which 
co-occur with the rise in Snowshoe Hare populations, 
are probably reinforced through density effects, where 
large numbers of hares are feeding in restricted habitats. 

For all parasites except Rauschia triangularis there 
were no significant (P = 0.05) differences between eco- 
regions or between sexes. Rauschia triangularis had a 
significantly higher (P < 0.027) abundance of parasites 
in males over females from the North Shore Eco-region. 
No other differences occurred for this parasite at the 
other eco-regions. We can offer no explanation for 
this. Erickson (1944) reports no significant difference 
between sexes for R. triangularis but does indicate that 
different areas may yield different prevalences for the 
parasite. He also found a high prevalence during late 
winter. 

An evaluation of the parasite standings based on 
weight increments for the entire population studied 
showed no significant differences (P 2 0.05) between 
weight classes in regards to Mosgovoyia pectinata and 
Rauschia triangularis (Table 4).Our results show that a 
significant majority (P < 0.04) of Obeliscoides cuniculi 
occur in the smaller hares. This differs from Keith et al. 
(1986), who report consistently higher values in adult 
hares during December-April (P < 0.001). As in our 
study, they did not show significant differences between 
sexes. Results similar to this study and Keith et al. 
(1986) are shown in Erickson (1944) and Keith et al. 
(1985). There is a tendency for R. triangularis to have 
a higher prevalence over O. cuniculi in the smaller 
hares, a situation similar to that reported by Keith et al. 
(1986) who reported a higher prevalences of R. trian- 
gularis (P = 0.059) among juveniles. Conversely, the 
larger hares contained significantly greater numbers 
(P < 0.001) of O. cuniculi over R. triangularis as is seen 
in Keith et al. (1986). 

An evaluation of multiple infections showed no ap- 
parent trends in the parasite burdens of males or fe- 
males. Sixty percent of the hares contained at least one 
species of parasite, 25.6 two, and 3.8 three; 10.3 percent 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


none. Dodds and Mackiewicz (1961) report greater par- 
asite diversities in their study. The trematode Dicro- 
coelium dentriticum and the nematode Trichostrongy- 
lus axei, which were reported from Emberly Island in 
Placentia Bay, are normally considered parasites of 
sheep and other ruminants. This island was used exten- 
sively for sheep grazing by residents and it appeared 
that the hares were paratenic (non specific or reservoir) 
hosts. Recent studies (in preparation) indicate the pres- 
ence and spread of new parasites within ungulates and 
insectivores which are attributed to the Coyote range 
expansion; however, no new parasites appear to have 
spread to the Snowshoe Hare from any of the recently 
(i.e., less than fifteen years) established non-native 
mammals. 


Acknowledgments 

We thank Mike McGrath, Senior Small Game and 
Furbearer Biologist, Inland Fish and Wildlife Divi- 
sion, Department of Tourism, Culture and Recreation, 
Government of Newfoundland and Labrador, St. 
John’s, Newfoundland, and the hunters and trappers 
who obtained the specimens utilized in this study. 


Documents Cited (marked * in text) 

Reynolds, J. J., B. K. Adams, and M. J. McGrath. 2004 
Status report on Snowshoe Hare population monitoring 
in insular Newfoundland, 1999 to 2004. Internal report 
to: The Government of Newfoundland and Labrador by 
the Department of Environment and Conservation, Wild- 
life Division. St. John’s, Newfoundland. 5 pages. 

Whitney, H. 2000 Common diseases of the Snowshoe Hare 
(1): Tapeworm cysts. Animal Health Division, Depart- 
ment of Forest Resources and Agrifoods, Government of 
Newfoundland and Labrador. St. John’s, Newfoundland. 
3 pages. 


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Dodds, D. G. 1957. Snowshoe hare research in Newfound- 
land. Department of Mines and Resources. Government 
of Newfoundland. St. John’s, Newfoundland. 7 pages. 

Dodds, D. G., and J. S. Mackiewicz. 1961. Some parasites 
and diseases of Snowshoe hares in Newfoundland. Jour- 
nal of Wildlife Management 25: 409-414. 

Erickson, A. B. 1944. Helminth infections in relation to 
population fluctuations in Snowshoe hares. Journal of Wild- 
life Management 8: 134-153. 

Gould, W. P., and W. O. Pruitt, Jr. 1969. First Newfound- 
land record of Peromyscus. Canadian Journal of Zoology 
47: 469. 

Heath, D. D. 1973. Resistance to Taenia pisiformis larvae 
in rabbits — I. Examination of the antigenically protective 
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Keith, L. B., J. R. Cary, T. M. Yuill, and I. M. Keith. 
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population. Journal of Wildlife Diseases 21: 233-253. 

Keith, I. M., L. B. Keith, and J. R. Cary. 1986. Parasitism 
in a declining population of snowshoe hares. Journal of 
Wildlife Diseases 22: 349-363. 


2004 


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Society of Parasitologists). Journal of Parasitology 68: 
131-133. 

Meades, W. J., and L. Moores. 1994. Forest site classifica- 
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Newfoundland. 2" edition FRDA Report 003, Forestry 
Canada, St. John’s, Newfoundland, Canada. 

Northcott, T. H., E. Mercer, and E. Menchenton. 1973. 
The Eastern Chipmunk, 7Zamias striatus, in Insular New- 
foundland. Canadian Field-Naturalist 88: 86. 

Parker, G. 1995. Eastern Coyote: The story of its success. 
Nimbus Publishing Ltd. Halifax, Nova Scotia, Canada. 
254 pages. 

Payne, R. F. 1976. Red squirrel introduction to Newfound- 
land. Canadian Field-Naturalist 90: 60-64. 

Rickard, M. D., and B. J. Coman. 1977. Studies on the fate of 
Taenia hydatigena and Taenia ovis larvae in rabbits, and 


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329 


cross immunity with 7aenia pisiformis larvae. Interna- 
tional Journal of Parasitology 7: 257-269. 

Scruton, D. A., K. D. Clarke, J. H. McCarthy, S. Forsey, 
D. M. Whitaker, G. I. McT. Cowan, E. Baggs, W. A. 
Montevecchi, J. M. Green, I. Bell, and L. J. Moores. 
1995. The Copper Lake Buffer Zone Study: Project site 
description and general study design. Canadian Techni- 
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2043: vi + 47 pages. 

Smith, F. R., and W. Threlfall. 1972. Helminths of some 
mammals from Newfoundland. American Midland Natural- 
ist 90: 215-218. 

Warren, G. L. 1970. The introduction of the masked shrew 
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Received 8 April 2002 
Accepted 24 August 2005 


Reconstructing Changes in Abundance of White-tailed Deer, Odocoileus 
virginianus, Moose, Alces alces, and Beaver, Castor canadensis, in 


Algonquin Park, Ontario, 1860-2004 


NorMAN W. S. QUINN! 


Algonquin Provincial Park, Box 219, Whitney, Ontario 
'Present address: R. R. #1, Bancroft, Ontario KOL 1CO Canada 


Quinn, Norman W. S. 2005. Reconstructing changes in abundance of White-tailed Deer, Odocoileus virginianus, Moose, 
Alces alces, and Beaver, Castor canadensis, in Algonquin Park, Ontario, 1860-2004. Canadian Field-Naturalist 
119(3): 330-342. : 


The history of White-tailed Deer, Odocoileus virginianus, Moose, Alces alces, and Beaver, Castor canadensis, in Algonquin 
Park since the 1860s is reviewed and placed in the context of changes to the forest, weather, and parasitic disease. Deer seem 
to have been abundant in the late 1800s and early 1900s whereas Moose were also common but less so than deer. Deer 
declined through the 1920s as Moose probably increased. Deer had recovered by the 1940s when Moose seem to have been 
scarce. The deer population declined again in the 1960s, suffered major mortality in the early 1970s, and has never 
recovered; deer are essentially absent from the present day Algonquin landscape in winter. Moose increased steadily 
following the decline of deer and have numbered around 3500 since the mid-1980s. Beaver were scarce in the Park in the 
late 1800s but recovered by 1910 and appear to have been abundant through the early 1900s and at high numbers through 
mid-century. The Beaver population has, however, declined sharply since the mid-1970s. These changes can best be explained 
by the history of change to the structure and composition of the Park's forests. After extensive fire and logging in the late 
1800s and early 1900s, the forest is now in an essentially mature state. Weather and parasitic disease, however, have also 
played a role. These three species form the prey base of Algonquin's Wolves, Canis lycaon, and the net decline of prey, 
especially deer, has important implications for the future of wolves in the Park. 


Key Words: Algonquin Park, Moose, Alces alces, Deer, Odocoileus virginianus, Beaver, Castor canadensis, Wolves, Canis 


lycaon, ticks, Dermacentor, Ontario. 


The biota of Algonquin Provincial Park, Ontario, 
have been under study since the mid-1930s and much 
information on fish and wildlife resources has 
accumulated since that time. There are, in addition, 
anecdotal records of the Park's major fauna from as 
early as the 1860s. There are particularly useful, if at 
times sporadic, data on White-tailed Deer (Odocoileus 
virginianus), Moose (Alces alces), and Beaver (Castor 
canadensis). Fluctuations in abundance of these species 
can be related to long-term change to structure and 
composition of the Park’s forests which, in qualitative 
terms at least, is well known (Strickland 1993). 
Recently, concern has been raised about the status of 
Wolves (Canis lycaon) in Algonquin Park (Theberge 
1998). Moose, White-tailed Deer, and Beaver collec- 
tively form the prey of Algonquin’s Wolves (Pimlott 
et al. 1969; Voigt et al. 1976; Forbes and Theberge 
1992; Forbes and Theberge 1996) and, ultimately, the 
fate of the Wolf population will depend on the status 
of these prey species. This paper is a review of the 
long-term history of Moose, White-tailed Deer, and 
Beaver in Algonquin Park. 


Methods 

The information presented in this paper from prior 
to about 1940 is essentially anecdotal. Post-1940, the 
data are from a mix of anecdotal references and popu- 
lation surveys, with formal surveys coming to dominate 


since about the 1960s. The “historic” information (pre- 
1940) is largely from land surveyors’ reports and annual 
reports of the Park Superintendent to the Minister 
responsible for Crown land. Post-1940 the data are from 
a variety of sources including annual reports of the 
Department of Lands and Forests (which became the 
Ministry of Natural Resources (O. M. N. R.) in 1972) 
which often featured Algonquin Park, unpublished 
reports from staff who worked at the Park's Wildlife 
Research Station, and, increasingly with time, published 
research papers. 

From about the mid-1950s the population surveys 
developed into standardized techniques accepted across 
the province. For Moose, this involved the gradual 
application of plot-based mid-winter aerial surveys 
that began in Ontario in about 1950 and have evolved 
into a formalized provincial survey technique (Bisset 
1996*). The currently accepted technique came into 
more or less its current form in the mid-1970s, but 
aerial survey data are reported here from 1950, the ear- 
lier surveys being somewhat less rigorous in design. 

Epizootics of Winter Tick, Dermacentor albipictus, 
can cause severe die-offs of Moose (Blyth and Hudson 
1987). In the early 1980s, E. Addison established a 
technique to assess the severity of tick infestation by 
modification of a “hair loss index” first developed in 
Alberta (Samuel and Barker 1979). The technique 
measures the extent of hair loss that is apparent on 


330 


2005 QUINN: WHITE-TAILED DEER, MOOSE, AND BEAVER IN ALGONQUIN PARK Aa 
* 
5 % - y- \) ' ome Hardwood-Pine Division 
at Beaver Survey 
et x (9 Markers 
\ AL . eo \ Beaver Survey 
| \ <2 Pp KY “ Flight-lines 
\ ‘i | 
\ | m ‘ ES Wilderness Zone 
= \ ¢ \ Ee Historic Deer Yards 
Le" ’ 
\ ae Major Water Areas 
10 0 10 Kilometers sh) : f r “)  Algonquin Park 
es Sees — Ce lewd Boundary 
\ \ f a 


FicurE |. Algonquin Park, showing the location of wilderness zones, historic deer yards, principal Beaver survey transects, 


and the east-west division of forest types. 


Moose in late winter. A survey is flown by helicopter 
on one day from 20-25 March in an approximately 
2000-km2 zone in the south-center of the Park to place 
at least 40 Moose in the five categories of hair loss 
proposed by Samuel and Barker. A simple arithmetic 
index, the Hair Loss Severity Index (H. S. I.), is then 
calculated by multiplying the number of Moose in 
each category by its severity rank (1-5), summing this, 
and dividing by the total Moose observed. The H. S. I. 
is used to predict the extent of mortality the follow- 
ing spring. 

Deer are much more difficult to census than Moose 
and have generally been surveyed by counting drop- 
pings or “pellet groups”, a technique that was first em- 
ployed in the U. S. in about 1940 (Bennet et al. 1940) 
and came into use in Ontario shortly afterwards. The 
technique was first applied in Algonquin Park in the 
mid-1950s and a protocol was developed that was 
used throughout south-central Ontario into the mid- 
1970s (Anonymous 1980*). These pellet group sur- 
veys provide the best reference to deer populations in 
the Park. As will be shown, the Park’s deer popula- 
tion declined drastically in the early 1970s and the 
technique, which is difficult in even ideal conditions 


and unreliable at low deer densities, was abandoned. 
Since the early 1970s deer surveys in the Park have 
consisted of late-winter aerial surveys of historic deer 
wintering areas (“yards”). Eight deer yards, some 
greater than 10 000 ha, were located across the Park 
(Figure 1) and supported high densities of deer (e.g., 
Stanfield 1957*) through mid-century. Several of these 
yards were surveyed in each of 1985, 1989, 1994, 
2000, 2001, and 2004. Every yard was flown at least 
once in the years listed and most were flown several 
times. The 2004 survey included several areas in the 
south-center of the Park that were outside the historic 
yards but with potentially good winter cover. The sur- 
veys were done in mid- to late March when snow is 
deepest. North-south transects were flown at | km inter- 
vals with fixed wing aircraft at approximately 140-km/h 
airspeed and deer tracks and trails tallied along the 
transects. In addition, deer sign was recorded during 
an aerial Wolf survey from 6-23 February 2002 (Pat- 
terson et al. 2004). Fifty-one hours were flown in a 
Bell 204 helicopter at approximately 100 m over 44 
5 x 5 km survey plots. Many of these plots were locat- 
ed in the historic yards and nearly all had at least some 
coniferous cover potentially suitable for deer. 


332 


Since 1955, Beaver have been surveyed in the Park 
by counting live colonies along aerial survey transects. 
The surveys have generally been done in late October 
after leaf fall but before ice-up when Beaver are active- 
ly storing food and evidence of fresh cutting (1.e., food 
piles) can be readily observed from the air. Results 
have been expressed as live colonies per unit area. 
There have been several different types of surveys and 
results of essentially all are reported here. The best 
data are from a long series of a survey first done in 1960 
by Rod Stanfield that consisted of two transects that 
each formed a circuit around the Park (a short segment 
of each flight was outside the Park, Figure 1). The 
survey was flown at a constant height of 800 feet by 
reference to points of elevation along the transects, 
and the observers line of sight established through a 
wing strut so that an observed band of 3200 feet was 
centered on the transect. Smith (1969*) summarized 
these data from 1960 — 1968. Stanfield described his 
survey technique, including the “way points” at the end 
of each transect, precisely, so that the survey can be 
repeated accurately. E. Addison reviewed and reinstated 
the survey in 1998 (Addison 1998*) and the survey was 
flown again in 1999 and 2003. 

Further information on specific survey methodolo- 
gies can be found in the papers cited above. Unpub- 
lished works and much of the original data for surveys 
referred to throughout this paper are found either in 
the O. M. N. R. Research Library in Peterborough, 
Ontario, or the Archives of Algonquin Park. 

Winter severity is a key factor in the productivity 
and survival of deer (e.g., Mech et al. 1987) and very 
severe winters can result in mortality of Moose (Bishop 
and Rausch 1974). Historic records of winter weather 
are presented and related to trends in abundance of deer. 
Monthly means of temperature were obtained from 
the Ontario Climate Center of Environment Canada in 
Downsview, Ontario, for Algonquin Park (Park Head- 
quarters at Cache Lake) from 1917 to 1972 and Dwight, 
Ontario, from 1973 to 2002 (the weather station was 
closed in the Park in 1973; Dwight is 20 km west of the 
Park boundary). Snow accumulation was obtained from 
a snow depth station that has been operating at the 
Park’s west gate since 1952. 2 

Forest management is also a key influence on all 
three species (e.g., Monthey 1984; Novak 1987; Peek 
1998). An overview of the history of logging and fire 
suppression in the Park is given in the next section 
before results of the various surveys are presented. 


Results 
History of Logging and Fire Suppression 

Algonquin Park is 7600 km? and is a multiple-use 
Park consisting of a complex array of zones in which 
varying degrees of activity are permitted. Complete 
protection is afforded to Wilderness Zones (Figure 1) 
but commercial logging is permitted in the Recreation- 
Utilization Zone, which comprises 78% of the Park. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


However, with various reserves of land for, for example, 
shoreline protection, only about % of the landbase is 
available for logging. This zoning system was estab- 
lished in the Master Plan of 1974 (Anonymous 1974*) 
and has been affirmed by Management Plans since then 
(Anoymous 1998*). 

The Park actually consists of two forests; the eastern 
third of the Park consists essentially of pine forests, pri- 
marily White Pine, Pinus strobus, with lesser amounts 
of Red Pine, Pinus resinosus, and isolated Jack Pine, 
Pinus banksiana, stands on well-drained, sandy outwash 
and rolling to flat terrain. The remaining two thirds, 
approximately 4600 km? of the Park’s west side, con- 
sists of tolerant hardwood forest: Sugar Maple, Acer 
saccharum, American Beech, Fagus grandifolia, Yellow 
Birch, Betula alleghaniensis, and Hemlock, Tsuga can- 
adensis, on glacial till over poorly-drained rugged 
terrain (Figure 1). 

As early as the 1830s (Strickland 1993) loggers 
entered the Park seeking White and Red pine “square 
timber” coincident with the removal of pine across 
northeastern North America at the time (Runkle 1985; 
Abrams and McCay 1996). Logging progressed rapidly 
up the main waterways of the Park; by 1866-67, 
30 000 pieces of square timber were removed annually 
(Strickland 1993). The square timber activity had peaked 
about 1864 and declined thereafter, the last square 
timber being cut in 1912 (Anonymous 2000*). White 
Pine, however, remained the focus of interest as saw- 
mills appeared before the square timber period ended. 

It seems that the removal of pine was extensive, 
nearly complete in places. Recent work analyzing rem- 
nant pine stumps (the fine remains of which can persist 
more than 120 years) suggests that 70 — 94% of the 
pine was removed from hardwood forests in the south- 
center of the Park (Simard 2001). The debris left from 
this logging fostered extensive fires primarily in the 
east-side pines (Fitzgerald 1890*, and see historic notes 
in Runge and Theberge 1974). Fires were frequent 
into the early 1900s although Superintendent G. W. 
Bartlett noted in several of his annual reports that these 
fires were “generally brought under control” (Bartlett 
1905-1921*). 

By the 1930s attention turned to hardwoods, in part 
because the pine had been depleted but also because 
of the development of new markets, including Yellow 
Birch veneers for construction of the Mosquito fighter- 
bomber of World War 2 (Anonymous 2000*). Remnant 
stands were often logged over for pulp (Runge and 
Theberge 1974). Fire apparently remained common 
until the early 1930s (Robinson 1933). 

Hard data on the extent of the removal of forest 
cover from all this activity is lacking but it appears that 
it was, in places at least, extensive. Photos exist from 
the late 1800s and early 1900s showing barren hillsides 
in the Park (Algonquin Park Museum Archives). White- 
tailed Deer, Moose, and Beaver are fundamentally 
adapted to early successional habitats (Novak 1987; 


2005 


Voigt et al. 1997; Peek 1998) and, from the point of view 
of food supply at least, habitat was probably favour- 
able for all three species through the first half of the 
1900s. Hall (1971), for example, noted that by 1893 
the logging of White Pine in Algonquin Park had been 
underway for 50 years and “considerable areas of the 
Park were undoubtedly in the secondary stages of forest 
succession that provide good beaver habitat”. Steph- 
enson and Hepburn (1958*) reported that continued 
logging since the 1800s had brought a “succession of 
areas” into a good productive state for deer. 

The harvest of Yellow Birch was gradually sup- 
planted by Sugar Maple, the primary use of which has 
shifted over time from sawlogs to pulp (Anonymous 
2000*). Hemlock, which is excellent winter cover for 
deer (Voigt et al. 1997), was not a priority species his- 
torically, nor is it today, but it was harvested in the 
late 1800s and early 1900s in the southwest of the Park 
for the tan bark industry (Strickland 1993) and then 
extensively in the 1960s for shoring timber, principally 
to build the Toronto subway (Wilton 1987). Wilton 
reported that 203 456 acres (82 339 ha) were cut over 
in six of the historic deer yards in the Park from 1952- 
1971. He pointed out that, although this created regen- 
eration for browse, “large volumes” of coniferous cover, 
particularly of Hemlock, were removed to the detri- 
ment of deer (Wilton 1987). M. Robinson made the 
same point in 1933, suggesting that “many of the for- 
mer yarding grounds have been destroyed by lumber- 
men removing the spruce, pine, and hemlock” (Robin- 
son 1933). 

As early as the late 1930s (Runge and Theberge 
1974) and definitely by the early 1950s (Anonymous 
2000*) diameter limit harvesting began to replace the 
more or less unregulated cutting. This evolved into par- 
tial cutting systems; the single-tree selection system, 
which was first employed in the late sixties in the hard- 
woods, and “uniform shelterwood” harvest in the pine 
stands on the east side. 

Selection cutting, which has been applied across the 
western 7 of the Park since the 1960s (Strickland 1993; 
Anonymous 2000*), retains a more or less intact forest 
canopy. Partial cutting systems can produce a consid- 
erable shrubby understory (Kelty and Nyland 1983) 
that is frequently renewed because stands are “treated” 
every 20-25 years. However, selective logging does not 
produce the biomass of browse that more aggressive 
logging systems do (e.g., Monthey 1984) and biologists 
in Ontario do not consider it of optimal benefit to 
browsers like deer (Voigt et al. 1997). Research on the 
effects of pine uniform shelterwood on ungulates is 
lacking but the system is normally effective at regen- 
erating White Pine (Anonymous 2000*) that grows 
to dominate the understory and thus may not produce 
large quantities of palatable browse. 

Forest fire is not normally active in mature tolerant, 
or “Northern”, hardwood forest found on the Park’s 
west side (e.g., Lorimer and Frelich 1994) and must 


QUINN: WHITE-TAILED DEER, MOOSE, AND BEAVER IN ALGONQUIN PARK 


3 oJ 


have become less frequent as the forest recovered. Fire, 
on the other hand, is naturally very frequent in the 
Park’s east side pines (Cwynar 1978). Fire was, at any 
rate, gradually eliminated from the landscape more or 
less coincident with the shift to low-impact logging. 
The introduction of fire towers and “fire ranging” air- 
craft in the late 1920s began a process that rapidly 
brought down the area of forest burned annually (Runge 
and Theberge 1974). Forest fire has been under essen- 
tially complete control in the Park for decades; al- 
though there are approximately two dozen lightning 
“starts” each year, fires rarely get past one hectare in 
size (Anonymous 2000*). 

Today, as a result of the elimination of fire and evo- 
lution to selection logging, the Park’s forests are in 
an essentially mature state. Seventy-seven percent of 
the Park’s hardwood zone is in mature tolerant hard- 
woods and the rest is in “static” wetland (e.g., Black 
Spruce, Picea mariana) or swamp hardwood forest, 
or mature intolerant hardwoods that are succeeding 
to Maple-Beech (Quinn 2004). The east-side pine for- 
ests are also primarily mature or uneven-aged (Anony- 
mous 2000*). The 22% of the Park that is in protected 
zones and has not been logged since at least the 1960s 
is also essentially mature forest, and, in places, ap- 
proaching “old growth” (as defined by Keddy 1994 and 
Tyrell and Crow 1994). 

In summary, the Park’s forests were extensively dis- 
turbed in the late 1800s and early 1900s but recovery 
began in the 1930s with the suppression of fire and 
adoption of low impact logging techniques. It is wide- 
ly accepted that the recovery of the Park’s forests has 
been detrimental to both White-tailed Deer (Runge 
and Theberge 1974; Wilton 1987) and Beaver. Moose 
have, as will be seen, increased through at least the later 
part of this period. 


Trends in Populations 
Deer 

The presettlement distribution of ungulates in central 
Canada is poorly understood, largely because bone 
deteriorates rapidly in the acidic soils of the Canadian 
Shield (Reid 1988). It is widely believed, however, that 
prior to European contact and the opening of the forests, 
deer did not range in Ontario north of approximately 
the location of Hwy 7, which is approximately 160 km 
south of the center of the Park (Matheson 1972; Smith 
and Borezon 1977; Smith and Verkruysse 1983). Deer 
bones were found in a dig of native hearths near Whit- 
son lake in Algonquin Park but could not be reliably 
dated and were presumed to be post-settlement (Burns 
19727). 

The earliest published evidence of deer in Algonquin 
Park is that of R. Bice in Wilton (1987) who reported 
that deer were found in the Park from at least the 1860s. 
There are a series of land survey records that suggest 
that White-tailed Deer (and Moose) were abundant in 
Algonquin through the late 1800s. Typically, the sur- 


334 


veyors noted that “the woods abound with moose and 
red deer” (Dickson 1883*; Byrne 1884*) or “red deer 
and moose roam all over the country” (Fitzgerald 
1890*). There is, however, one dissenting voice in 
1887; “little, if any, game was seen during the survey” 
(White Township.) (Fitzgerald 1887*). 

From 1905 to 1921, G. W. Bartlett, Park Superin- 
tendent, wrote annual reports on Algonquin Park to 
the Minister of Lands and Forests. These reports con- 
sistently suggest that deer were very abundant. Phrases 
like “the deer are here simply in thousands and ... in- 
creasing” are found throughout (e.g., Bartlett 1907*). 
In 1911 Bartlett reported that “deer are so abundant 
they can be seen from the hotel veranda in numbers” 
and in 1910 that deer supported great hunting on the 
Park’s boundaries and that “the Park is a great feeder 
for the surrounding country”. 

Robinson (1933) reported that deer numbered in 
the tens of thousands in 1921 but had declined to 
“possibly not more than three thousand” by 1933. 
Although there are no snow depth records for the time, 
winter temperatures were not unusually cold, indeed 
rather warm for the late 1920s — early 1930s (Figure 
2). Robinson in fact emphasized deteriorating habitat, 
in part because of the suppression of fire, as the cause 
of the decline (Robinson 1933). The population, how- 
ever, apparently recovered because C. H. D. Clarke 
reported a “chronic overpopulation” of deer in 1945 
and reports an estimate of 13 deer/square mile, from a 
very early pellet group survey, a number he considered 
“suspiciously low” (Clarke 1945*). A. Leopold also 
listed the Algonquin deer population as among those 
“overpopulated” in North America at the time (Leo- 
pold et al. 1947). 

Population estimates from pellet group surveys 
began to appear regularly in the 1950s. Stephenson 
(1958*), for example, estimated 53 deer/square mile 
in a wintering yard (where, of course, deer were con- 
centrated) in Biggar Township in 1957. He was re- 
searching means to ameliorate “heavy cropping” of 
Yellow Birch by deer at the time (Anonymous 1957*, 
see research section page 72) so deer must have been 
abundant. Estimates from pellet group surveys for 
the area around Swan Lake, in the southwest corner 
of the Park suggested 15 deer/square mile in winter 
and 12 in summer in 1957 (Stephenson and Hepburn 
1958*). Estimates of 12 deer per square mile would 
have put the Park population at approximately 36 000 
animals at the time. 

There was apparently significant winter mortality 
(a “die-off”’) of deer in the winters of 1958-1959 and 
1959-1960 (Runge and Theberge 1974). These winters 
were indeed cold and snow accumulation was excep- 
tional (Figure 2). The range was also overbrowsed at 
the time; Grant Taylor, retired Park naturalist, recalls 
that all regeneration from ground to five feet in height 
was browsed (G. Taylor, personal communication). The 
losses, however, may not have been extensive because 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


there is an estimate of 69 deer/square mile in a yard 
in the Kiosk area in 1961 (Anonymous 1961*) and deer 
are reported to have “recovered” by 1965 (Anonymous 
1965*). B. Stephenson reported 10 deer/mile? in sum- 
mer within a 3 square mile study area located in the 
south of the Park in 1960 (Pimlott et al. 1969, page 
29). Deer must have been common in the Park in the 
1960s because the late Roy Anderson reported hay- 
ing “hundreds” of road-killed deer to dissect for his 
studies of meningeal worm, Parelaphostrongylus ten- 
uis, in the 1960s (R. Anderson, University of Guelph, 
personal communication). 

Notwithstanding this, deer apparently declined grad- 
ually through the 1960s (Rutter 1964; Runge and 
Theberge 1974) and this culminated in a severe die-off 
in the winters of 1970-1971 and 1971-1972. Wilton 
(1970*) estimated a population of 8090 in 1970 but 
that had declined to 2800 by 1972 (Wilton and Trodd 
1972*). The decline was also documented in King 
(1976) who reported extensive mortality of deer in cen- 
tral Ontario in the winter of 1970-1971 and a density 
of only 2.02 deer/square mile in Pembroke District 
(which included Algonquin Park) in spring of 1972. 
King attributed the decline to winter severity, and snow 
depths were indeed exceptional (Figure. 2). Snow was 
also persistent those two winters; the peak of snow 
depth in Algonquin Park is normally the second week 
in March but it was mid-April in 1971 and late March 
ren NOPE. 

The sharp decline of deer that occurred in Algon- 
quin in the early 1970s happened throughout central 
Ontario (King 1976). Deer populations have since re- 
covered around the Park but the Algonquin popula- 
tion never recovered. Deer are fairly common in the 
Park in summer; Park staff have been conducting road 
counts since 1999 and one deer is seen per approxi- 
mately 270 km driven on Hwy 60 in May (roughly 
one deer per eight Moose, unpublished data: Algonquin 
Park files). However, deer are very scarce in winter. 
Essentially no deer or deer tracks were seen in any of 
the aerial deer yard surveys from 1984-2004 described 
earlier (including the 2002 Wolf survey). The only 
exception to this is the yards in the Southern “pan- 
handle” (Figure 1) which still support deer. Deer or 
deer tracks are also rarely observed during winter aerial 
Moose surveys of the Park. 


Moose 

Moose were almost certainly on the early Algonquin 
landscape. Moose are known to have existed in Maine 
in the 1600s and in Quebec at the same latitude as 
Algonquin Park in the 1500s (Reeves and McCabe 
1998). Furthermore, Peterson (1955) suggests that 
Algonquin Park was within the range of Moose in 1875. 
Intriguingly, however, it was noted in the Royal Com- 
mission Report on the founding of the Park in 1893 
that Moose were unknown to the Indians prior to 
1870 when they made their first appearance “at least 


2005 QUINN: WHITE-TAILED DEER, MOOSE, AND BEAVER IN ALGONQUIN PARK 335 


Maximum Snow Depth, Algonquin Park 1953 - 2002 


100 


DEPTH (CM) 


20 
1950 1960 1970 1980 1990 2000 
YEAR 


Mean Monthly Minimum Temperature, Jan. - March, Algonquin Park, 1918 - 2002 
-8 


-10 
-12 
O 
o -14 
2 
oF 
w -16 
a 
= 
uJ 
fun 
-18 
-20 
-22 
1910 1930 1950 1970 1990 2010 


FIGURE 2. Long term trends of snow depth and winter temperature in Algonquin Park. Arrows point to years of deer die-offs. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Algonquin Park Moose Population - 1950-2003 


336 
5000 
4000 
LUI §=3000 
” 
O 
O 
= 
© 2000 
a 
1000 


“die off" years 


ba 


1955 


1965 


1975 1985 1995 2005 


YEAR 


FIGURE 3. Moose population estimates, Algonquin Park, 1950 — 2003. 


in recent times” after crossing over from Quebec 
(Anonymous 1893*). 

Moose were, at any rate, well established in the Park 
by the late 1800s as evidenced by reports of the early 
land surveyors (e.g., Dickson 1883*; Byrne 1884*; 
Fitzgerald 1890*) discussed earlier. Reports of waste- 
ful killing of Moose by hunters in the late 1800s in 
the area soon to be Algonquin Park (Anonymous 
1893*) also suggest high numbers. Bartlett's annual 
reports, however, suggest that Moose were less com- 
mon than deer in the early 1900s (Bartlett 1905-1921*). 
Moose are consistently referred to as “increasing” or 
“greatly increased” but as secondary to deer in these 
reports. Moose may, however, have been locally num- 
erous as “in great numbers along the Nipissing River” 
in 1908 (Bartlett 1908). 

There is a gap in reports of Moose abundance 
through the 1920s and 1930s, although Robinson 
(1933) reported that Moose “have increased greatly 
in numbers” after the “disappearance of the deer” he 
had noted from the early 1920s; the implication is that 
Moose were scarce relative to deer up to about 1921 
which is in agreement with Bartlett. It seems, however, 
that Moose had declined again by the 1940s. Robb 
(1942) reports seeing only 19 Moose (but 254 deer) 
in Beaver surveys in the Park in 1939-1940 and C. H. 
D. Clarke suggested that Moose were very scarce in 
1945. Clarke reported that although Moose are “dis- 


tributed across the Park” only “one specimen found 
dead’”’could be examined and that “Algonquin Park is 
hardly the place to study moose in Ontario” (Clarke 
1945*). He further reported that only two Moose pellet 
groups were found in 98 deer pellet group plots. 

Population estimates for Moose post-1945 are shown 
in Figure 3 and show a more or less steady increase 
until the present. Estimates in Figure 3 post-1985 are 
directly from the Park’s data files; those prior to 85 are 
from DeVos (1952*); Pimlott et al. (1969, page 32); 
Wilton and Pashuk (1983*) (to which the 1984 and 
1985 estimates were appended); and several Depart- 
ment of Lands and Forests annual reports from 1954- 
1968. The population estimates from 1956-1958 are 
extrapolations from estimates of Moose density for 
Pembroke District (which included Algonquin Park). 
Also, as few as 12 plots were used in some of these 
early surveys so they may not have been very accurate. 
Years missing in the inventory post-1974 are the result 
of poor survey conditions. 

The late March survey of winter tick induced hair 
loss was done 14 times from 1984 to 2004. The mean 
H. S. I. was 1.90 (range 1.18 — 3.48) and there was 
evidence of heavy losses of Moose in spring following 
three of five surveys (March of 1989, 1992 and 1999) 
in which the index was > 1.95. Moose carcasses, or the 
stench thereof, were frequently reported by the public 
throughout spring of 1989 and 1992 (personal obser- 


QUINN: WHITE-TAILED DEER, MOOSE, AND BEAVER IN ALGONQUIN PARK 


a7 


Algonquin Park Beaver Population - 1940 - 2003 


2005 
0.9 
0.8 
0.7 

= 

4 

lu 0.6 

cc 

S 

eee 

a 

whe 

a 

2 

5 0.3 

= 

ay 
0.1 
0.0 

1930 1940 1950 1960 


1990 


1970 1980 2000 


YEAR 


FiGuRE 4. Beaver population estimates, Algonquin Park, 1940 — 2003. 


vation). In 1999 (while I was on leave) the H. S. I. 
reached its highest and there was evidence of extensive 
mortalilty (B. Sandilands, Park Planner, personal com- 
munication). Unfortunately, aerial Moose surveys in 
two of the three winters following this apparent heavy 
mortality (1990 and 2000) were cancelled or only partly 
completed and thus the population effects are unclear 
[and note that the survey in winter of 1993 actually 
showed an increase (Figure 3)]. 


Beaver 

There is little about Beaver from the early land 
surveys except a reference from 1890 about scattered 
Beaver meadows “now more or less dried up” (in 
White Township) (Fitzgerald 1890*). James Dickson 
Provincial Land Surveyor, however, wrote in 1888 that 
trapping had greatly reduced the Beaver population 
in the region soon to become Algonquin Park and 
Peter Thompson, first Park Superintendent, reported 
that “scarcely a beaver could be seen” (Hall 1971). By 
1899, however, the Beaver population had reportedly 
recover-ed and were so numerous by 1909 that the Park 
Superintendent recommended that trapping (which had 
been suspended) be reinstated (Hall 1971). Superin- 
tendent Bartlett's reports suggest that Beaver flour- 
ished from 1905-1921. In 1906, for example, Bartlett 
wrote that “fur bearing animals have greatly increased, 
especially the beaver’. Bartlett reported in 1908 that 


“the beaver can be found in numbers upon every lake, 
river, pond and creek” and in 1909, “in large healthy 
colonies wherever there is a drainage ditch” (he lament- 
ed the extent of “nuisance” problems Beaver were caus- 
ing in the 1908 report). The remaining reports frequently 
note Beaver increasing, for example “the annual in- 
crease of which (Beaver) runs into the thousands” in 
LO: 

Information is lacking for the Park for the 1920s. 
However, Beaver were apparently plentiful across the 
province up until 1923 but declined sharply (province- 
wide) in 1924 and remained low, and of concern, until 
1930 (Anonymous 1923-1930*). No reason is given 
for the decline and it is not clear if the decline occurred 
also in Algonquin Park. 

Robb (1942) reported a density of 0.29 Beaver 
colonies/km? in 1940 in the south-center of the Park 
but Hall (1971) believed that Robb underestimated the 
population and reported that Beaver populations had 
fluctuated between 0.40 — 0.80 colonies/km* from 
1955 to 1971. Beaver must have been relatively abun- 
dant in the 1950s because R. Stanfield reported an 
“unusually high density” in 1955 and between 58 and 
77 colonies/100 “bodies of water” in 1957 and that this 
was high “relative to all other Districts except North 
Bay” (Stanfield 1957*). 

Population density estimates of Beaver (colonies/ 
km’) for the Park from 1940 to 2003 are shown in 


338 


Figure. 4. These data are primarily Stanfield’s survey 
[summarized in Smith (1969*)] and its recent contin- 
uation (Addison 1998*), but include several other inde- 
pendent estimates (Robb 1942; Wilton 1974*; Regan 
1978*). The data suggest that the Beaver population 
has declined sharply since the late 1970s (Figure 4). 
This is reflected in independent studies in the south- 
center of the Park by J. Fryxell, who reported a signi- 
ficant 50% decline of Beaver from 1987-1998 (Fryxell 
2001). 


Discussion 

In summary, White-tailed Deer appear to have been 
common in Algonquin Park in the late 1800s while 
Moose were present but secondary to deer. Deer re- 
mained abundant and apparently increased through 
the early 1900s until a decline from the early 1920s to 
the early 1930s. Deer had recovered by the early 1940s 
and were common through the 1950s but began to 
decline in the 1960s and suffered severe losses due to 
winter severity in the early 1970s and have not recov- 
ered. Moose were apparently secondary to deer in the 
early 1900s and probably relatively scarce until the 
decline of deer in the 1920s when they are reported to 
have increased. By the 1940s, however, Moose had 
declined (coincident, apparently, with the recovery of 
deer) and appear to have been quite rare by the mid- 
1940s. Aerial surveys show an increase in the Moose 
population beginning in the mid-1950s that accelerated 
with the decline of deer in the early seventies. Moose 
are now the dominant ungulate in what was a Wolf- 
deer system through most of the last century. Beaver 
were scarce until the formation of the Park in 1893 
whereupon they increased rapidly and were very com- 
mon through the early 1900s. There are little data on 
Beaver until aerial surveys first began in 1940. Surveys 
and qualitative assessments suggest that Beaver were 
abundant and more or less stable at about 0.5 col- 
onies / km? through mid-century but began to decline 
in the mid-1970s and are now at less than half of their 
peak abundance. 

The conventional explanation for the observed 
trends in White-tailed Deer and Moose is that the re- 
moval of coniferous cover and gradual maturation of 
the forest have been detrimental to deer and, conse- 
quently, of benefit to Moose (Robinson 1933; Runge 
and Theberge 1974; Wilton 1987). Presumably, mat- 
uration of the forest also accounts for the decline of 
Beaver (Addison 1998*). The system, however, is more 
complex than just that and a more detailed discussion 
is in order. 

The opposition of Moose and deer numbers over 
time is intriguing given the potential virulence of 
meningeal worm to Moose (Anderson 1964). Menin- 
geal worm is widely distributed in eastern North 
America and present in Algonquin Park and one is 
tempted to conclude that Moose increased in the Park 
as deer, and the rate of transmission of P. tenuis, de- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


clined. Wildlife managers in the Park and elsewhere 
have assumed this effect occurs, at least with relatively 
high deer densities (e.g, Karns 1967). However, the 
ecological role of meningeal worm is poorly under- 
stood and evidence for the degree of its impact on 
Moose has been challenged (Nudds 1990). Neverthe- 
less, the parasite must remain implicated as having a 
potential role in the recurring divergence of Moose and 
deer numbers in the Park. 

There are other enigmatic questions regarding Moose 
and White-tailed Deer. For example, the scarcity of 
deer in the Park in winter while deer are abundant all 
around the Park is hard to explain. It is, as discussed, 
widely believed that this is a result of the deterioration 
of habitat in the Park but also because winter weather 
is more severe than for the surrounding landscape. The 
latter is true; for example mean monthly minimum 
temperatures in winter are lower in a zone centered 
on the core of the Park than the surrounding area 
(Anonymous 1984). Yet deer winter in large numbers 
in yards almost adjacent to the Park boundary where 
winter weather is only marginally different. For exam- 
ple, deer winter in the hundreds in yards near Dorset 
(J. MacDonald, O. M. N. R., Bracebridge, personal 
communication) only 15 km from the Park boundary 
and at 1100 m elevation vs 1250 m for Hogan Lake, one 
of the original yards in the Parks’s core. The structure 
and composition of the forest around Dorset and the 
surrounding landscape, is, superficially at least, similar 
to that of the Park (personal observation). Why, then, 
are deer so scarce in the Park in winter? 

Wolves may be the answer to this question. We oc- 
casionally find Wolf-killed deer in the Park in mid- 
winter in places where their presence was not previ- 
ously obvious. In the 2002 aerial Wolf survey one of — 
only four deer observed was a recent Wolf kill anda | 
second was being pursued by Wolves. It seems that 
Wolves are very effective at finding the few deer that do | 
winter in the Park (Forbes and Theberge 1996). Pos- 
sibly, White-tailed Deer have not been able to reoccupy | 
the Park after the die-offs of 1971 and 1972 because | 
Wolves have essentially excluded them (at least in win- 
ter). There is in fact a body of evidence that predation 
is disproportionately heavy on isolated groups of deer 
wintering away from the core of large yards (Kolen- 
osky 1972; Nelson and Mech 1981; Patterson and 
Messier 2000). 


2005 


winter in hereditary yards even where quality of habitat 
is poor (Voigt et al. 1997). Possibly, supplementary 
feeding has, over time, created a fixed “culture” of 
seasonal movement out of the Park. Lewis and Rong- 
stad (1998) showed that supplementary feeding of 
deer can influ-ence migration. 

One hypothesis, then, to explain the recent history of 
deer in Algonquin Park is as follows: The decline of 
deer in central Ontario in the early 1970s was partic- 
ularly severe in Algonquin Park. Deer recovered around 
the Park but were slower to do so in Algonquin because 
of poorer quality of habitat and somewhat more diffi- 
cult winters. Supplementary feeding, which began in 
earnest shortly after the decline of deer, presented a 
draw that developed into fixed winter migrations and 
the few deer that try to break that mode are killed by 
Wolves. 

Moose also present a dilemma. With the exception of 
some island populations in northwestern Ontario, 
Algonquin Park has supported the highest Moose pop- 
ulation density in the Province for years; typically 
showing densities three times higher than in northern 
management units (O. M. N. R., Wildlife Surveys and 
Records). The Park, however, is almost the antithesis 
of good Moose habitat. Quality Moose habitat 1s tra- 

_ ditionally viewed as landscapes that are extensively 
disturbed; boreal forests with a mix of burns and/or 
clearcuts amidst winter cover and aquatic feeding areas 
(e.g., Peek 1998). Algonquin Park, or at least its western 
two thirds where Moose densities are highest, is a 
closed-canopy tolerant hardwood forest with very few 
openings larger than a few hundred square meters 
(Anonymous 2000*; Quinn 2004). Fire has not been 
active on the Algonquin landscape for decades and, 
even when not suppressed, is very rarely a stand des- 
tructive event in tolerant hardwoods (Lorimer and 
Frelich 1994). Partial cutting systems in hardwoods 
can provide substantive browse (Kelty and Nyland 
1983) but much less than clearcuts (Monthey 1984). 
Why, then, are there so many Moose in the Park’? 
Here, again, the answer may lie in predation for, on 
the Park’s west side at least, there is no effective pred- 
ator acting on Moose. Hunting by a local aboriginal 
community is permitted on the east side of the Park 
in a zone corresponding roughly to the extent of pine 
forest (about 40% of the Park’s area). The west side 
(hardwood) area is, however, unhunted. Further, Al- 
} gonquin’s Wolves, although capable of killing Moose, 
| are principally a “deer-eating” type (Pimlott et al. 
1969), and primarily a scavenger of Moose in winter 
| (Forbes and Theberge 1992, 1996). A hypothesis 
jj regarding the Algonquin Moose population thus goes 
ij as follows: Moose are at high densities in Algonquin 
¢|| Park because of the lack of an effective predator, not 
{| the quality of habitat. Habitat is marginal but accept- 
able; selective cutting provides a limited but contin- 
uous supply of browse and coniferous cover, while 
i) generally unsuitable for deer, is adequate for Moose. 
Relatively low predation pressure, and very low den- 


QUINN: WHITE-TAILED DEER, MOOSE, AND BEAVER IN ALGONQUIN PARK 


eS) 
eS) 
\Oo 


sities of deer (and thus P. tenuis) allow Moose to do 
well in a forest environment that is not optimal. 

A less well understood player in this system is the 
Black Bear (Ursus americanus). Black Bears prey on 
Moose calves and can limit Moose populations (Stewart 
et al. 1985). Data on bear predation on Moose calves 
in Algonquin, however, are sparse. Moose are believed 
to calve preferentially on islands and peninsulas in the 
Park to avoid bears (Addison et al. 1990) but Garner 
(1994) showed a predation rate of bears on Moose 
calves in the Park of only 8% which is low relative to 
that suggested by removal studies of Brown Bears 
(Ursus arctos) in Alaska (Miller and Ballard 1992) 
and Black Bears in Saskatchewan (Stewart et al. 1985). 

The Algonquin ecosystem, and particularly the com- 
plex interplay of Wolf-prey, habitat, weather, and para- 
sites will continue to evolve and present challenges to 
Park managers. The future of the Park’s Wolves will 
depend largely on trends of these prey populations. This 
basic truism has largely been overlooked in the dis- 
cussion on the status of Algonquin’s Wolves which 
has focused on human killing around the Park as the 
cause of the possible decline (Theberge 1998). This 
paper suggests that declining prey may be at least as 
important a factor. Algonquin’s Wolves are small and 
prey preferentially on deer (Pimlott et al. 1969; Forbes 
and Theberge 1996). The near absence of deer in winter 
and decline of Beaver, an important summer food, 
must be stressful to the population, as has been docu- 
mented in Minnesota (Van Ballenberghe and Mech 
1975). The increase of Moose has in part compensated 
but Algonquin’s wolves are primarily scavengers of 
Moose in winter (Forbes and Theberge 1996) and may 
find food energy from Moose readily available only in 
winters of tick induced mortality. 

Algonquin Park, and its wolves and prey, do not 
exist in isolation. Indeed, recent work suggests that 
Algonquin’s wolves are not unique but genetically 
identical to and freely interbreeding with wolves around 
the Park (Grewal 2001). The larger population, a dis- 
tinct taxon, the “Eastern Canadian Wolf’, Canis lycaon, 
(Wilson et al. 2000) extends from Manitoba to Quebec 
and numbers approximately 10 000 (Van ZylI de Jong 
1996*; White et al. 2001*). External phenomena that 
are both minor and local, like feeding deer, and great 
and global, like climate change, will perhaps have as 
much influence on future trends in the distribution and 
abundance of C. /ycaon as changes within the Park 
itself. 


Acknowledgments 

A great many biologists and other Park staff, far 
too numerous to mention, contributed to the collection 
of the data that have been presented. Particular recog- 
nition should, however, be given to those that have 
worked in the Park in the more recent past and in par- 
ticular, M. Wilton, E. Addison, R. Anderson, D. Voigt, 
G. Forbes, J. Theberge, B. Stephenson, and R. Stanfield. 
E. Hovinga has directed Moose surveys in the Park in 


340 


recent years and is one of a long list of Moose flyers 
with strong stomachs and large bladders. V. Michalsen 
assisted with the figures. B. Patterson provided useful 
comments on an early draft. 


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Received 29 May 2003 
Accepted 15 November 2004 


The Distribution and Habitat Selection of Introduced Eastern Grey 
Squirrels, Sciurus carolinensis, in British Columbia 


EMILY K. GONZALES 


Centre for Applied Conservation Research, University of British Columbia, Forest Sciences Centre, 3004-2424 Main Mall, 


Vancouver, British Columbia V6T 174 Canada 


Gonzales, Emily K. 2005. The distribution and habitat selection of introduced Eastern Grey Squirrels, Sciurus carolinensis, in 
British Columbia. Canadian Field-Naturalist 119(3): 343-350. 


Eastern Grey Squirrels were first introduced to Vancouver in the Lower Mainland of British Columbia in 1909. A separate 
introduction to Metchosin in the Victoria region occurred in 1966. I surveyed the distribution and habitat selection of East- 
ern Grey Squirrels in both locales. Eastern Grey Squirrels spread throughout both regions over a period of 30 years and 
were found predominantly in residential land types. Some natural features and habitats, such as mountains, large bodies of 
water, and coniferous forests, have acted as barriers to expansion for Eastern Grey Squirrels. Given that urbanization is 
replacing conifer forests throughout southern British Columbia, it is predicted that Eastern Grey Squirrels will continue to 


spread as habitat barriers are removed. 


Key Words: Eastern Grey Squirrels, Sciurus carolinensis, distribution, habitat selection, invasive, British Columbia. 


The vast majority of introduced species do not suc- 
cessfully establish populations in novel environments 
(Williamson 1996). Many successful non-native species 
are human comensals which thrive in human-modified 
environments (Williamson and Fitter 1996; Sax and 
Brown 2000). Eastern Grey Squirrels (Sciurus caroli- 
nensis; hereafter EGS), a charismatic species endemic 
to eastern and central North America, have been inten- 
tionally introduced to Great Britain, Italy, Ireland, 
South Africa, Australia, and western North America 
(Robinson and McTaggart-Cowan 1954; Seebeck 1984; 
Gurnell 1987; Lever 1994). Their gregarious nature has 
made them a popular addition to city parks and they 
generally flourish in this land type whether they are 
native or introduced. The ability of EGS to establish 
populations in association with humans is likely a key 
to their success as an invader. This study aims to update 
what is known about the spread and distribution of 
introduced EGS in southwestern British Columbia. I 
also analysed their habitat usage and predicted that EGS 
would be found predominantly in residential areas, areas 
that more closely resemble their native environment. 

EGS occur naturally in the eastern United States 
and central Canada. Their distribution coincides with 
eastern hardwood forests; especially nut producing trees 
such as oaks (Quercus spp.), hickories (Carya spp.), 
and, formerly, American Chestnut (Castanea dentata). 
Their coats are often grey or black and, more rarely, 
red, blonde, and white. EGS eat nuts, fruits, fungus, 
and, opportunistically, eggs and nestlings (Orr 1971). 
EGS are most successful in deciduous forests, mixed 
deciduous-conifer forests, and residential areas (Bark- 
alow and Shorten 1973; Pasitschniak-Arts and Ben- 
dell 1990; Riege 1991). The highest densities of EGS 
in the wild occur in mature maple-oak forests and the 
lowest in fir-cedar forests (Riege 1991). Very high den- 
sities of EGS have been observed in residential areas 


such as backyards, parks, and cemeteries (Pasitschniak- 
Arts and Bendell 1990). EGS co-occur with North 
American Red Squirrels (Tamiasciurus hudsonicus) 
throughout much of their range where habitat special- 
ization and not competition determines the differences 
in their distributions (Riege 1991). 

EGS have had negative economic and ecological 
effects in the places where they have been introduced. 
In Europe, EGS strip the bark from trees, damaging 
both native and plantation species (Kenward and Parish 
1986). EGS may also replace European Red Squir- 
rels (Sciurus vulgaris) in Great Britain (e.g., Lloyd 
1983; Wauters and Gurnell 1999; Gurnell et al. 2004), 
although studies demonstrating the mechanism of 
replacement have been inconclusive. Attempts to erad- 
icate, control, or slow the spread of EGS have been 
costly failures, in part due to a lack of understanding 
of the population biology and methods of spread of 
EGS (Sheail 1999; Gonzales 2000*; Bertolino and Gen- 
ovesi 2001). Conifer forests and plantations, mountain- 
ous terrain, and bodies of water have slowed spread or 
were barriers to movement, while deciduous forests 
and poplar plantations have provided suitable habitat 
for EGS in Europe (Williamson and Brown 1986; 
Wauters et al. 1997; O Teangana et al. 2000a, 2000b). 
EGS were found in conifer forests in Great Britain, but 
the maintenance of those populations required immi- 
gration from neighbouring habitat types (Kenward and 
Hodder 1998; Bryce et al. 2002). The role of conifer 
forests in limiting the spread and establishment of 
EGS has led to broad scale land management recom- 
mendations that involve the maintenance of conifer 
forests/plantations and the removal of large seed decid- 
uous trees, particularly oaks (Kenward and Hodder 
1998; Bryce et al. 2002). 

Conifer forests historically dominated British Co- 
lumbia and habitat conversion from conifer forest to 


343 


344 


residential development with deciduous trees may have 
increased the amount of suitable habitat for EGS. 
Today, residential areas in the Victoria region and 
Lower Mainland are well treed with deciduous species. 
Gardens, bird feeders, garbage and hand-outs pro- 
vide an abundance of food while trees and human 
dwellings provide nesting locations. 

The first introduction of EGS to British Columbia 
occurred in 1909 when at least six individuals from 
the New York Park Department were brought to the 
peninsula of Stanley Park in Vancouver (Steele 1993). 
The population increased and had achieved a stable size 
by the 1920s (Robinson and McTaggart-Cowan 1954). 
Robinson and McTaggart-Cowan (1954) surveyed the 
population and natural history characteristics of EGS 
in Stanley Park in 1950. EGS were found predominant- 
ly in deciduous forests, mixed deciduous-conifer 
forests, and developed areas rather than in the conifer 
forests that dominate Stanley Park. Their diet was pri- 
marily composed of hand-outs from visitors, nuts from 
horticultural trees, and samaras from native maple trees. 
Although EGS are known to eat eggs and nestlings, 
these authors did not find any occurrences of this. The 
authors speculated that EGS would remain confined 
to Stanley Park because it was effectively an island, 
surrounded by the Burrard Inlet and downtown Van- 
couver. EGS remained on the peninsula until the mid- 
1970s, when populations began spreading to adjacent 
areas (Merilees 1986, 1992; Gonzales 1999). 

EGS were first introduced to Vancouver Island in 
1945 (Ringuette 2004). A small population was intro- 
duced to Beacon Hill Park in the city of Victoria. They 
disappeared, however, and the current population of 
EGS are believed to be descendants of an introduction 
that occurred in the autumn of 1966. Three individuals 
were acquired from southwestern Ontario for a private 
game farm in Metchosin, within the Greater Victoria 
area (Guiguet 1975; Fraser 1987). These EGS were 
released some years after their arrival and had spread 
to neighbouring municipalities by 1975 (Guiguet 1975). 


Study Areas 

Both study sites are in southwestern British Colum- 
bia, Canada, and are separated from each other by a 
40 km ocean strait (Figure 1). Both sites have mild cli- 
mates and mean annual temperatures of 8°C-10°C. The 
Lower Mainland, an area of approximately 1500 km?, 
is located in the moist Coastal Western Hemlock bio- 
geoclimatic zone (Pojar et al. 1991). Prior to devel- 
opment, this region was predominantly conifer forest. 
Natural areas are still dominated by conifers, particu- 
larly Western Hemlock (T'suga heterophylla), Western 
Red Cedar (Thuja plicata), and coastal Douglas-fir 
(Pseudotsuga menziesii). Residential areas have num- 
erous horticultural trees, both in private gardens and 
along boulevards. For example, the Vancouver Park 
Board maintains over 124 000 trees comprising over 
600 species and cultivars, the majority of which are 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


deciduous (Vancouver Board of Parks and Recreation 
2005"): 

The Victoria region is an area of approximately 
460 km? located in the Coastal Douglas-fir biogeocli- 
matic zone (Nuszdorfer et al. 1991) on the southeastern 
tip of Vancouver Island. Natural areas are dominated 
by coastal Douglas-fir, Shore Pine (Pinus contorta), 
and Western Red Cedar. Residential areas have a large 
variety of horticultural trees. The most common boule- 
vard trees are Japanese flowering cherries (Prunus spp.) 
and Horse Chestnuts (Aesculus hippocastanum). Garry 
Oaks (Quercus garryana) are endemic to the region 
and the acorns may provide a food source for EGS. 

Given the negative effects of EGS in Europe, there 
is speculation that EGS have negative ecological im- 
pacts in British Columbia. EGS may eat and damage 
a sufficient number of Garry Oak acorns to impede 
their regeneration (Bruemmer et al. 2000*). EGS may 
also compete for resources with endemic squirrels 
(Garry Oak Ecosystem Recovery Team 2003). North 
American Red Squirrels are endemic to the Victoria 
region and Douglas Squirrels (Jamiasciurus douglasii) 
and Northern Flying Squirrels (Glaucomys sabrinicus) 
are endemic to the Lower Mainland. 


Methods 

I conducted field surveys and solicited sightings of 
squirrels from the public, particularly naturalists, from 
1996 to 2000, through paper and telephone surveys 
and a web page. A series of questions determined the 
presence or absence, location, coat colour and the year 
that EGS were first sighted at a location. A lack of 
squirrel sightings from the public surveys could be 
due to an absence of squirrels or a sampling effect. To 
ensure representative sampling across all land types, 
I conducted surveys in areas that were underrepresent- 
ed by the public surveys. Using Geographic Informa- 
tion Systems (GIS), I overlaid a grid comprised of 
5 km x 5 km cells onto each study area. Cells without 
sightings of squirrels were identified by land type on 


Lower Mainland 


50 P=) 0 
| we 


50 Kilometers 


UTM: 475000, 5425000 Victoria 


Region 


FiGuRE |: Map of southwestern British Columbia showing 
the Lower Mainland and the Victoria region. 


2004 


a digital landscape map and the total area of each land 
type was computed. I calculated the number of public 
responses by land type to get a measure of “survey 
effort” by land type. Surveys were conducted in each 
of the land types so that each land type had an equal 
level of survey effort proportional to its presence on 
the landscape. Trained volunteers and I conducted 
175 hours of auditory and visual searches for EGS, 
Douglas and Red squirrels. Surveys occurred from 8 am 
to 11 am, when squirrels are generally more active. 
Surveyors stayed at a point location for 10 minutes 
before moving to the next location approximately 
100 m away. In addition to watching for EGS, Dou- 
glas, or Red squirrels, surveyors also listened for their 
calls to confirm their presence. 

I also acquired submission records for EGS, Dou- 
glas and Northern Flying squirrels from three wildlife 
shelters located in different municipalities in the Lower 
Mainland for the years 1983 to 2003. Submission 
records begin in 1983, before EGS had spread to most 
municipalities in the Lower Mainland. The shelter 
records were used to augment the public surveys to plot 
the spread of EGS by year across the Lower Mainland. 
The records did not state the specific location where 
the squirrels were found, and therefore were not used 
in the habitat selection analysis. 

I tested whether EGS use the land types proportional 
to their availability using a log-likelihood chi-square 
test with 95% confidence limits and a Bonferroni cor- 
rection (Neu et al. 1974; Manly et al. 1993). The loca- 


500 


@ Eastern grey 
| CO Douglas 
| » Northern flying 


400 ; 


300 


200 


# of squirrels 


100 


GONZALES: EASTERN GREY SQUIRRELS IN BRISTISH COLUMBIA 


345 


tions of EGS, given by nearest street intersection, 
landmark or GPS coordinates, were converted into 
UTM coordinates and overlaid onto digital maps using 
ArcView 3.2 (Environmental Systems Research Insti- 
tute 1998*). Six land type categories, each ground- 
truthed with field surveys, were categorized on maps 
and included: (a) Agriculture (farmland and treeless 
fields); (b) Parks (municipal, regional, and provincial 
parks); (c) Schools (including fields, treed and unde- 
veloped areas around a building or buildings; (d) Indus- 
trial (city centres and heavily developed areas); (e) 
Residential (urban and suburban areas); and (f) Open 
(cleared areas). Department of National Defence lands 
in the Victoria region (fields with some development) 
were included in the Open category. Land types in 
which squirrels were never sighted, such as lakes and 
gravel pits, were excluded from the analysis. 


Results 

In total, 212 responses were received for the Lower 
Mainland and 383 for the Victoria region. Wildlife 
shelter records provided an additional 4937 municipal 
locations of squirrels for the Lower Mainland. Over the 
20 years, there were a total of 264 Northern Flying 
Squirrels, 656 Douglas Squirrels, and 4017 EGS 
brought to the wildlife shelters (Figure 2). EGS first 
arrived at the shelters in 1985 and their numbers have 
increased exponentially until 2000. EGS did not move 
contiguously through the municipalities in the two 
study sites. While it is likely that there were gaps in 


1993 1998 2003 


Year 


FIGURE 2. Number of Eastern Grey, Douglas and Northern Flying squirrels from wildlife shelters in the Lower Mainland 


from 1983 to 2003. 


346 


the temporal data, the translocation of EGS by humans 
also explains the discontiguous pattern of expansion. 
The surveys revealed that residents and pest control 
companies translocated live EGS, often beyond a “bar- 
rier” such as a river or mountain in the hopes that 
nuisance EGS would not return (Pynn 1999*). By 
2004, EGS occurred throughout the Lower Mainland 
(Figure 3) and the Victoria region (Figure 4). 

The habitat selection analysis revealed that EGS did 
not use the land types in proportion to their prevalence 
(x? << 0.001, df = 5). EGS were found predominantly 
in the land type when the proportion of EGS is signif- 
icantly above the proportion of the land type’s avail- 
ability (Figures 5 and 6). EGS in the Lower Mainland 
were found predominantly in Residential land types 
and not in Agricultural or Open land types (Figure 5). 
EGS in the Victoria region were also found predomi- 
nately in Residential land types and not in Agricul- 
tural, Open, or Parks land types (Figure 6). 


Lion's Ba 


West 
Vancouver 


University 
Endowment 
Lands 


a: a~ 


Richmond i 


me? 
zd 
ub Vy 


Presence of EGS by Year 
mee No data available 


Cette 1909-1969 
r] 1970-1979 
{fT} 1980-1984 


1985-1989 
1990-1994 


[ee | 

i ‘: 
MM 1995-1999 ee 
ay 


2000-2004 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Discussion 

EGS have spread predominantly toward Residential 
areas. In the Lower Mainland, the Pacific Ocean to the 
west and conifer forest and mountains to the north 
have acted as barriers to their spread. Suitable habi- 
tat, specifically residential development, is available 
south and east and spread has proceeded further and 
more quickly in these directions. A similar pattern was 
seen in the Victoria region where EGS spread through- 
out the eastern peninsula of Vancouver Island, toward 
residential areas rather than west and north into areas 
dominated by conifer forests. The results suggest that 
EGS may continue to invade British Columbia, but are 
more likely to do so in residential areas and are un- 
likely to spread in areas still dominated by conifer 
forests. The land type categories were very general, 
however, because of the size of the region. I used the 
most detailed landscape data available for the breadth 
of the study areas. Localized studies that defined spe- 


N 
Hai Moody 
/ , Pitt Meadows 
/ Coquitlam ys 


/ 


White Rock _ 


Langley 


8 Kilometers 


FIGURE 3. The spread of Eastern Grey Squirrels in the Lower Mainland from 1909 to 2004. 


2004 GONZALES: EASTERN GREY SQUIRRELS IN BRISTISH COLUMBIA 347 


fi 
| 


North Saanic ull) 


Presence of EGS by Year il 
5] 1970-1974 Witt} ih Central Saenich 
oo 1975-1979 Lt 


1 980- 1 984 Highlands 
1985-1989 
TI] 1990-1994 


san Saanich mee 


athretyseieiese Fee 
sy yey ete sys 


* £48 $22 ctona 


1995-2000 “a 
poy ). i ~/ 
| 
aw, it ls of 
sets Bk Bay 
ee cS ood . 


+t 
44% 


pean 
Metchosin Penis 


2 0 2 4 Kilometers 
bal aan 


Beh shy) 
wateuexeceret 
A “SS AL 


FiGurE 4. The spread of Eastern Grey Squirrels in the Victoria Region from 1970 to 2000. 


Lower Mainland 


80% 
@ Land Type 
0 
2 60% O Eastern Grey Squirrels 
Wd 
oO. 
en 40% 
o 
— 
o 
en | 
20% 
0% 


Agriculture Parks Residential Schools Commercial Open 


Land Types 


FiGuRE 5. Habitat selection of Eastern Grey Squirrels in the Lower Mainland. The proportion of Eastern Grey Squirrels 
found in each land type with 95% confidence intervals (4? << 0.001, d.f. = 5). Eastern Grey Squirrels were found 
predominantly in Residential land types and not in Agricultural or Open land types. 


348 


cies of trees, the locations of bird feeders and other 
resources, would further our understanding of the 
specific habitat needs of EGS at fine scales. 

While some introduced species can have profoundly 
negative effects on native species, the majority are 
benign (Williamson 1996). Further, landscape conver- 
sion that favours many successful invaders is simultane- 
ously detrimental to native species. The result may 
appear to be the displacement of native species by non- 
native species, but this interpretation 1s confounded 
by the changes in the landscape. The squirrels endemic 
to the Lower Mainland evolved in a landscape dominat- 
ed by conifer trees. If there has been a decline in 
endemic squirrels, it is more likely that the decline is 
related to habitat loss than to direct competition with 
EGS. If EGS competitively displaced native squirrels in 
the Lower Mainland, we would expect to see the extir- 
pation of Northern Flying Squirrels and Douglas Squir- 
rels where EGS were first introduced. All three species 
of squirrels have been present at the original site of the 
introduction, Stanley Park, for nearly 100 years (per- 
sonal communication, Michael Macintosh, Supervisor 
at the Vancouver Board of Parks and Recreation). 

Although EGS and North American Red Squirrels 
co-occur throughout most of their native range through 
habitat segregation (Riege 1991), North American Red 
Squirrels are thought to be declining due to EGS in 
the Victoria region (Bruemmer et al. 2000*; Garry Oak 
Ecosystem Recovery Team 2003). The decline of Red 
Squirrels, however, may be coincident with or exacer- 
bated by habitat loss. For example, a survey respondent 
reported that North American Red Squirrels disap- 
peared from southeastern municipalities in the Victoria 
region in the 1940s as residential development in- 
creased. The replacement of natural areas with residen- 
tial development allows EGS to fill an “empty niche” 
rather than actively displace native squirrels. EGS may 
negatively impact native squirrels on local scales where 
their habitats overlap such as mixed deciduous-conifer 
forests or rural residential areas, but at a regional scale, 
EGS are likely to co-occur as long as there 1s suffi- 
cient suitable habitat for each of them. 

There is some concern that EGS reduce the regen- 
eration of Garry Oaks, an important species in en- 
dangered Garry Oak ecosystems, by eating acorns, 
notching acorns, and stripping the bark from young 
trees (Bruemmer et al. 2000*; Garry Oak Ecosystem 
Recovery Team 2003). This belief, however, 1s con- 
tradictory to the literature that supports a net benefit 
for tree regeneration from scatter hoarding squirrels 
(e.g., Vander Wall 2001; Goheen and Swihart 2003). 
For example, North American Red Squirrels have re- 
cently expanded their range in Indiana, areas previous- 
ly dominated by EGS. EGS are concurrently declining 
due to land conversion, an interesting contrast to EGS 
in British Columbia. Goheen and Swihart (2003) pre- 
dict that the replacement of scatter hoarding EGS 
with larder hoarding Red Squirrels will result in the 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


decline of Black Walnut (Juglans nigra), Red Oak 
(Quercus rubra), and Bur Oak (Quercus macrocarpa) 
regeneration. 


Conclusions 

A disproportionate number of successful introduc- 
tions of non-native species occur in disturbed habitats 
(Williamson 1996). As humans convert natural areas 
into residential areas, they facilitate the spread of non- 
native, human comensal species such as EGS. Starting 
with just a few individuals, EGS have spread through- 
out the Lower Mainland and Victoria region. After an 
initial lag time following introduction, EGS took 
approximately 30 years to spread throughout each of 
the study areas. They have also been translocated to 
communities beyond but discontiguous from the two 
study areas. EGS are found significantly more often in 
residential areas than other land types. While EGS are 
rarely found in conifer forests, the general trend to 
replace conifer forest with residential development will 
facilitate their spread into areas that have previously 
acted as barriers to EGS distribution. 

Some residents in British Columbia have observed 
the spread of EGS in the Lower Mainland and Victoria 
region with trepidation because introduced species can 
have negative impacts on native species. EGS are 
believed to impede the regeneration of Garry Oak and 
competitively displace native squirrels in both study 
areas. Neither of these hypotheses has been tested and 
it is possible that EGS have negative ecological impacts 
in British Columbia. There are reasons, however, to be 
optimistic. EGS are beneficial to hardwood tree regen- 
eration, particularly oaks, in regions where EGS are 
native. EGS co-occur with North American Red Squir- 
rels over large parts of North America through differ- 
ences in habitat selection, so competitive displacement 
of Red Squirrels by EGS on Vancouver Island would 
not be expected to occur. Further, Northern Flying 
Squirrels, Douglas Squirrels, and EGS have continued 
to be present in Stanley Park for almost 100 years. 

Regardless of whether EGS have a negative impact 
on native species in British Columbia, they are consid- 
ered undesirable as a non-native species. In Europe, 
broad scale land management such as the maintenance 
of conifer woodlots rather than hardwoods has been 
recommended to limit populations of EGS (Kenward 
and Hodder 1998; Kenward et al. 1998; Bryce et al. 
2002). The maintenance of conifer forests will also 
limit the spread of EGS in British Columbia. Education 
should also be a key component because translocations 
of EGS by humans have facilitated their spread across 
barriers such as bodies of water, mountains, and conifer 
forests. 


Acknowledgments 

This project required data over large study areas and 
I am thankful to the many sources that contributed data. 
Digital maps were made available through the Ministry 


2004 


GONZALES: EASTERN GREY SQUIRRELS IN BRISTISH COLUMBIA 


349 


Victoria Region 


80% 


60% 


40% 


Land Type/EGS 


20% 


0% 


Agriculture Parks 


Residential 


@ Land Type 
O Eastern Grey Squirrels 


Schools Commercial Open 


Land Types 


FiGureE 6. Habitat selection of Eastern Grey Squirrels in the Victoria region. The proportion of Eastern Grey Squirrels found 
in each land type with 95% confidence intervals (x << 0.001, d.f. = 5). Eastern Grey Squirrels in the Victoria region 
were found predominately in Residential land types and not in Agricultural, Open, or Parks land types. 


of Environment, Lands and Parks through the depart- 
ments of the Crown Land Registry Services, Geograph- 
ic Data BC, and the Strategic Planning Department of 
the Greater Vancouver Regional District. Undergraduate 
volunteers at Simon Fraser University and the Uni- 
versity of Guelph were instrumental in data collection, 
field surveys, and data entry. The members of the Van- 
couver Natural History Society, the Burke Mountain 
Natural History Society, the White Rock Natural His- 
tory Society, and the Victoria Natural History Society 
provided many of the squirrel locations. I appreciate the 
comments of Tom Nudds, Jane Reid, Simone Runyan, 
Peter Arcese, Isaac McEachern, and two reviewers who 
significantly improved earlier versions of the manu- 
script. 


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Received 25 June 2003 
Accepted 29 August 2005 


The Fathead Minnow, Pimephales promelas, in New Brunswick 


DALE J. Hoop! and RUDOLPH FE. STOCEK? 


'DJ Hood & Associates Ltd., 90 Crystal Drive, Waasis, New Brunswick E3B 9B8 Canada 
270 Baxter Court, Fredericton, New Brunswick E3B 6M1 Canada 


Hood, Dale J., and Rudolph F. Stocek. 2005. The Fathead Minnow, Pimephales promelas, in New Brunswick. Canadian Field- 


Naturalist 119(3): 351-354. 


A second record for the occurrence of the Fathead Minnow (Pimephales promelas) in New Brunswick, the first for the south- 
west portion of the province, is reported. This is the first documented occurrence of the species in the province in 44 years, 
and the first indication that the Fathead Minnow occurs outside the upper Saint John River system in New Brunswick. 


Key Words: Fathead Minnow, Pimephales promelas, distribution, occurrence, habitat, St. Croix River, New Brunswick. 


The Fathead Minnow (Pimephales promelas) is 
widely distributed in Canada (Scott and Crossman 
1973), occurring from New Brunswick in the east, west 
through southern Quebec, all of Ontario, and large parts 
of Manitoba, Saskatchewan, Alberta, and a small area 
in the southern Northwest Territories. Various checklists 
and documents on fish species in New Brunswick iden- 
tify the Fathead Minnow as occurring in the province 
(Scott and Crossman 1959, 1973; Gorham 1970s). 

The status of the Fathead Minnow in New Brunswick 
has been uncertain, and sometimes a subject of debate, 
since it was first recorded here. Scott and Crossman 
(1959) reported the first record of the Fathead Minnow 
in New Brunswick waters. That record was based on 
the collection of two specimens in 1958 from an un- 
named tributary to the Saint John River near Ed- 
mundston (Royal Ontario Museum Catalogue Num- 
ber 19705): 

Here, we report the collection of Fathead Minnows 
in southwestern New Brunswick during 2002. Based 
on the new records, it is important to alert researchers 
and fisheries resource managers of the potential for the 
species to be more widely distributed in the province 
than previously believed. 

The Fathead Minnow is characteristically a small 
schooling species found in freshwater and brackish 
environments. The habitats associated with the species 
vary greatly throughout its range (Scott and Crossman 
1973). It is found in lakes, ponds, and streams of vary- 
ing sizes, ditches, reservoirs, and residual pools of 
intermittent streams, usually in sluggish or still water 
with a muddy bottom and abundant floating and sub- 
merged vegetation. The species is tolerant of high tem- 
perature, turbidity, low oxygen concentration, and high 
salinity. 

Fathead Minnows were found at three sites in the 
watershed of a small unnamed tributary to the St. Croix 
River at St. Stephen, while conducting systematic elec- 
trofishing spot-checks for fish presence on | and 3 Au- 
gust 2002. All sites were located in settings that rep- 
resented areas highly disturbed by human activities. 


Table | summarizes details of the settings and habitat 
characteristics at sites of the historical and new records. 

The sites where Fathead Minnows were collected 
in 2002 are approximately 250 km from the location 
where they were collected in 1958 by Scott and Cross- 
man (1959). Figure 1 shows the localities where Fathead 
Minnows have been documented in New Brunswick. 

The Fathead Minnow was the only species of fish 
present at the three sites where it was collected in 2002. 
These sites are located upstream of Queensway Street 
(St. Stephen), in a section of stream with an average 
slope of 1.7%. A fourth site, sampled downstream of 
Queensway Street in a section of stream with an aver- 
age slope of 7.2%, had a fish species assemblage that 
consisted of American Eel (Anguilla rostrata), Brook 
Trout (Salvelinus fontinalis), Creek Chub (Semotilus 
atromaculatus) and Lake Chub (Couesius plumbeus). 
The culvert under Queensway Street appears to be a 
barrier to upstream fish passage. 

Six specimens of Fathead Minnows were captured 
at SC1, nine specimens at SC2, and one specimen at 
SC3. Fork length was measured for specimens at SC] 
(range = 34-56 mm; mean = 40 mm) and SC2 (range 
= 32-61 mm; mean = 43 mm). One specimen retained 
from SC1 and two specimens from SC2 were deposit- 
ed in the collections of the New Brunswick Museum 
(NBM), catalogued as NBM 1181 and NBM 1182, 
respectively. All other specimens were released back 
into the waters from which they were captured. 

These new records raise questions related to the spot- 
ty distribution of populations of the Fathead Minnow 
in the province: are the recorded populations native or 
have they resulted from introductions of the species 
in some localities? Scott and Crossman (1973) iden- 
tified the waters of western New Brunswick as being 
on the northeast fringe of the distribution range for 
the Fathead Minnow. 

The Fathead Minnow has been recorded in approx- 
imately 100 lakes and ponds in the neighbouring State 
of Maine, occurring in all regions of the state (David 
Halliwell, Maine Department of Environmental Pro- 


351 


Vol. 119 


THE CANADIAN FIELD-NATURALIST 


352 


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2005 


Hoop and STOCEK: FATHEAD MINNOW IN NEW BRUNSWICK 


United States of America 


353 


20 40 60 80 100km 
KILOMETRES 


FiGureE |. Distribution of the Fathead Minnow in New Brunswick. Locality designations correspond to those in Table 1. 


tection, personal communication to DJH; Richard Jor- 
dan, Maine Department of Inland Fisheries and Wild- 
life, personal communication to DJH). Several of these 
locations occur in watersheds that drain to the Maine— 
New Brunswick boundary waters of the Saint John 
River or the St. Croix River systems. The waters in 
Maine where the Fathead Minnow has been recorded, 
nearest the sites in St. Stephen, are Upper and Lower 
Mud Lakes (Machias River watershed) and Orie Lake 


(St. Croix River watershed), located approximately 
23 km southwest and 49 km northwest, respectively. 
The records of occurrence of the Fathead Minnow 
in Maine are biased by the sampling effort associated 
with management of lakes that support important re- 
creational fisheries for species of salmonids and small- 
mouth bass (Micropterus dolomieu). If sampling was 
expanded to other waters, it is believed that the Fat- 
head Minnow would be found at other locations in 


354 


Maine (Richard Jordan, Maine Department of Inland 
Fisheries and Wildlife, personal communication to 
DJH). 

Whittier et al. (2000) described the general distri- 
bution and native status of 24 minnow species in north- 
eastern USA. They determined the Fathead Minnow to 
be native to the lakes where they collected it in central 
and northern Maine. However, the native status of this 
species has not been determined for the various loca- 
tions where it has been recorded in the waters of east- 
ern Maine (Richard Jordan, Maine Department of In- 
land Fisheries and Wildlife, personal communication 
to DJH). The Fathead Minnow is considered represen- 
tative of intrastate introduced native fish species in 
Maine, as a result of illegal baitfish transplants (Hal- 
liwell 2003). 

Live baitfish can be transported long distances. Con- 
sequently, bait-bucket transfer has been suggested as 
a major vector for illegal introductions and the occur- 
rence of species in areas outside their native ranges 
(Litvak and Mandrak 1993, 2000; Rahel 2000). The 
use of baitfish for recreational fishing is widespread 
in Maine and the Fathead Minnow is a species com- 
monly used (Richard Jordan, Maine Department of 
Inland Fisheries and Wildlife, personal communica- 
tion to DJH). In New Brunswick, the use of baitfish 
is limited by provisions in the provincial angling reg- 
ulations. The use or possession of live fish as bait 1s pro- 
hibited, except on tidal waters and on boundary waters 
between Maine and New Brunswick. On boundary 
waters, the only live fish that can be used or possessed 
for bait are fish taken from the waters being fished. 

Due to the proximity of the St. Stephen sites to boun- 
dary waters where the Fathead Minnow could be used 
as a baitfish, its occurrence at these sites could be the 
result of a bait-bucket transfer. However, this is not 
certain with the information available. 

In order to confirm the status of the 2002 records, the 
Atlantic Canada Conservation Data Centre (ACCDC) 
and various museums in eastern Canada were contact- 
ed to determine if they hold unreported data or col- 
lections of the Fathead Minnow from New Brunswick. 
The ACCDC reported that there were no records of the 
Fathead Minnow in New Brunswick other than that 
reported by Scott and Crossman in 1959 (Kate Bredin, 
ACCDC, personal communication to DJH). Consul- 
tation with representatives from Canadian and major 
regional museums confirmed that they hold no unre- 
ported collections of the Fathead Minnow from the 
province (Sylvie Laframboise, Canadian Museum of 
Nature, personal communication to DJH; Donald 
McAlpine, New Brunswick Museum, personal com- 
munication to DJH; John Gilhen, Nova Scotia Muse- 
um of Natural History, personal communication to 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


DJH; Erling Holm, Royal Ontario Museum, personal 
communication to DJH). 

The lack of additional occurrence records for the Fat- 
head Minnow in New Brunswick might be a reflection 
of the actual distribution of the species in the province, 
or perhaps a lack of effort among some fisheries field 
personnel to distinguish between the numerous species 
of minnows encountered. Historically, individual species 
of minnows have been given little attention in fisheries 
surveys, often being lumped together into the generic 
category of “minnows” (Whittier et al. 2000). 

The new records in 2002 indicate the potential for 
the species to be more widely distributed in the prov- 
ince than previously believed. It is unknown if the 
populations of the Fathead Minnow at St. Stephen are 
native to southwestern New Brunswick, or if they have 
been introduced. 


Acknowledgments 

We express our thanks to Adam Carr for assistance 
in the field. Donald McAlpine, Curator of Zoology, 
New Brunswick Museum, verified identification of the 
specimens collected during 2002. Erling Holm, Assis- 
tant Curator of Fishes, Royal Ontario Museum provided 
verification of identification of specimens catalogued 
as ROM 19705. Dan McDonald, Dillon Consulting 
Limited, prepared the location map. 


Literature Cited 

Gorham, S. W. 1970. Distributional checklist of the fishes of 
New Brunswick. The New Brunswick Museum. 32 pages. 

Halliwell, D. B. 2003. Introduced fish in Maine. Maine Aquat- 
ic Biodiversity Project series: Focus on Freshwater Bio- 
diversity. 12 pages. 

Litvak, M. K., and N. E. Mandrak. 1993. Ecology of fresh- 
water baitfish use in Canada and the United States. Fish- 
eries 18(12): 6-13. 

Litvak, M. K., and N. E. Mandrak. 2000. Baitfish trade as 
a vector of aquatic introductions. Pages 163-179 in Non- 
indigenous Freshwater Organisms: Vectors, Biology, and 
Impacts. Edited by R. Claudi and J. H. Leach. Lewis Pub- 
lishers. CRC Press. 464 pages. 

Rahel, F. J. 2000. Homogenization of fish faunas across the 
United States. Science 288: 854-856. 

Scott, W. B., and E. J. Crossman. 1959. The freshwater fishes 
of New Brunswick: a checklist with distributional notes. 
Contributions of the Royal Ontario Museum, Division of 
Zoology and Palaeontology Number 51. 37 pages. 

Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes 
of Canada. Fisheries Research Board of Canada Bulletin 
184. 966 pages. 

Whittier, T. R., D. B. Halliwell, and R. A. Daniels. 2000. 
Distributions of lake fishes in the northeast — II: the min- 
nows (Cyprinidae). Northeastern Naturalist 7: 131-156. 


Received 16 July 2003 
Accepted | June 2005 


Vision and its Relationship to Novel Behaviour in St. Lawrence River 
Greenland Sharks, Somniosus microcephalus 


Curis J. HARVEY-CLARK!, JEFFREY J. GALLANT’, and JOHN H. Batt? 


‘Animal Care Center, University of British Columbia, 6199 South Campus Road, Vancouver, British Columbia V6T 1W5 
Canada, email: chclark @interchange.ubc.ca 

?Aqualog Society, P.O. Box 483, Drummondville, Quebec J2C 6W3 Canada, email: aqualog @aqulaog.ca 

3Aquatron Laboratory, Department of Oceanography, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada; e-mail: 
John.Batt@ Dal.ca 


Harvey-Clark, Chris J., Jeffrey J. Gallant, and John H. Batt. 2005. Vision and its relationship to novel behaviour in St. Lawrence 
River Greenland Sharks, Somniosus microcephalus. Canadian Field-Naturalist 119(3): 355-359. 


Rarely observed Greenland Sharks, Somniosus microcephalus, were recorded at shallow depths by divers employing underwater 
video in the St. Lawrence River, in association with a seasonal concentration of Capelin (Mallotus villosus) in May-June 2003. 
We recorded unique proximity-induced display motor patterns in these sharks, which have not been recorded in underwater 
observations of Arctic Greenland Sharks. Arctic sharks have a high incidence of blindness due to an ocular copepod parasite, 
Ommatokoita elongata. The absence of parasite-induced blindness in St. Lawrence Greenland Sharks, in contrast to endemic 
blindness in the Arctic population, may allow sharks in this region to more readily visually recognize the presence of conspecifics 
and potential prey. Improved visual acuity may therefore allow St. Lawrence River sharks to express a different behavioural 
repertoire than Arctic sharks, with resulting changes in intra- and inter-specific aggression and predatory behaviour. 


Key Words: Chondricthyes, Squaliformes, Somnosidae, Greenland Shark, Somniosus microcephalus, display behaviour, copepod, 


Ommatokoita elongata. 


The Greenland Shark, Somniosus microcephalus 
(Bloch and Schneider 1801) has seldom been ob- 
served under natural conditions, and until this report, 
had been observed by divers underwater only rarely 
in Arctic waters (Caloyianis 1998). This giant member 
of the elasmobranch Order Squaliformes is a slow 
moving, primarily deep water shark reaching at least 
6.4 metres in length (Compagno 1984), found in cir- 
cumpolar regions and in cold, deep water to depths 
of at least 2200 metres (Herendorff and Berra 1995). 
Greenland Shark stomachs contain a wide range of 
prey including marine invertebrates, fish and mammals 
as well as terrestrial vertebrates such as Caribou Rangi- 
fer tarandus (Compagno 1984), and the role of this spe- 
cies as a predator, or primarily a scavenger, of marine 
mammals is controversial (Bigelow and Schroeder 
1953; Ridoux et al. 1998; Lucas and McAlpine 2002). 

St. Lawrence River Greenland Sharks free of the 
ocular copepod parasite Ommatokoita elongate were 
observed by divers during an unusual inshore migra- 
tion associated with seasonal concentrations of Cape- 
lin Mallotus villosus. Here we report on closely ap- 
proached sharks that exhibited display motor patterns 
not previously described in this species. 


Methods 

We employed SCUBA and underwater video in the 
region of Baie-Comeau, Quebec, Canada (49°16.9'N, 
68°07.2'W, Figure 1), to record Greenland Shark size, 
sex, physical condition, swimming speed and behav- 


iour in May and June 2003. The occurrence in in- 
shore waters of this elusive species was accompanied 
by an unusually high seasonal inshore concentration 
of Capelin (Mallotus villosus) massing around docks 
and bottom structures which was the highest recorded 
in the region over the previous five years (F. Gregoire, 
Canadian Department of Fisheries and Oceans, Institute 
Maurice Lamontaigne, Mont-Joli, Quebec, personal 
communication 2003). Both juvenile Harbour Seal 
(Phoca vitulina) and Grey Seal (Halichoerus grypus), 
potential prey species of the sharks, were locally abun- 
dant and were observed during fieldwork and diving 
activities. 

Observations were made between 09:00 and 17:30 
Eastern Standard Time and occurred in seawater depths 
from 10.0 to 28.0 m (mean=18.1 m +/- 5.8 m 95% con- 
fidence interval, CI). Seawater temperatures ranged 
from 1.0°C to 9.0°C (mean= 4.4°C 4/- 3.1°C 95% CI). 
We employed a standard report form and interviewed 
local professional divers, who first reported the pres- 
ence of sharks in the region. 

Videotape of sharks was analyzed for size scale using 
adjacent objects of known maximum size (Plumose 
Anemones, Metridium senile, maximum height of 
40 cm) and visible time code (hours/minutes/seconds) 
to determine swimming speed. We recorded 68.2 video 
minutes during prolonged encounters with four sharks 
over a three-day period and documented six other en- 
counters occurring between 26 May and 12 June 2003 
in the same region. 


305 


356 


BAIE-COMEAU @ 


GULF OF 
ST. LAWRENCE 


FiGurE |. Location of Greenland Shark sightings in the St. 
Lawrence River in the vicinity of Baie-Comeau, Prov- 
ince of Quebec. 


Results 

Of the 10 shark encounters, the range in estimated 
length was from 2.5 to 4.5 m total length (mean = 
2.8 m +/- 0.34 m CI 95%). In four sharks sex was 
determined (3 female, | male), with one markedly 
“girthy” 3.0 m female shark believed to be gravid. In 
one instance three sharks were seen separately on the 
same dive and in one instance two sharks were ob- 
served simultaneously within 5 meters of each other on 
diverging courses. Shark swimming speeds ranged 
from 0.10 m/s to peak at 1.40 m/s (mean = 0.62 m/s 
+/- 0.31 m/s 95% Cl). Greenland Sharks were capable 
of rapid acceleration from 0.10 m/s to 1.40 m/s and 
could outpace divers when swimming in a straight 
line, were highly manoeuvrable and were capable of 
changing depth and direction rapidly. In one case, the 
same 2.50 m female shark identified by scar patterns 
was seen repeatedly over a 3-day period in the same 
location, in each case within 50 m of the same posi- 
tion when encountered. Female sharks were noted to 
have more scarring than males, generally linear scar- 
ring on the tail, pectoral fins and dorsum of the cau- 
dal peduncle. We postulate that these scars may have 
resulted from nuptial or combat related behaviour, as 
observed in other shark species (Compagno 1984). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


FIGURE 2. The rostrum of a female Greenland Shark illus- 
trating typical loss of pigmentation seen anterior and 
adjacent to prominent nares. The cornea lacking 
attached parasitic copepods, and the dorsal opercu- 
lum are clearly visible. 


Both sexes exhibited a whitish cruciate pattern which 
appears to be a loss of pigmentation on the anterior 
rostrum, and which may be the result of bottom for- 
aging activities using the rostrum (Figure 2). A male 
shark was noted to have a single left clasper, although 
this species generally possesses two claspers (Com- 
pagno 1984). 

During encounters sharks were found swimming 
approximately 1.0 to 2.0 meters above the substrate 
towards divers on converging courses, and following 
close approach swam on sinusoidal courses, in some 
cases performing complete or half circle manoeuvres 
over a wide radius (20 to 30 m). These circling man- 
oeuvres resulted in sharks maintaining continuous eye 
contact with divers, and in two instances sharks left 
the bottom to ascend upwards at an acute angle and 
closely approach a diver above them in mid-water, 
apparently in investigative fashion. 

When sharks were closely approached from the 
lateral or anterior aspect (2.0 m or less from the eye), 
they initially displayed a motor pattern which includ- 
ed rapid bilateral ventral deflection of the pectoral 
fins (Figure 3 a, b), deceleration, and adoption of a 


FiGuRE 3. Female Greeland shark showing (a) normal pectoral fin position and (b) marked ventral deflection of pectoral fins 
as part of a motor pattern observed consistently when closely approached by divers. 


2005 


head-down attitude with the front of the animal 0.1- 
0.3 m above the substrate, the trunk slanted upward and 
the tail held higher off the bottom, the dorsum curved 
upward, the axis of the body being held approximately 
10 degrees off horizontal, and the mouth held slightly 
open (Figure 4). This posture was maintained from one 
to several seconds, after which the shark would accel- 
erate and swim away from the diver or change course 
to avoid the diver. This motor display was repeatedly 
exhibited by the four sharks we observed when close- 
ly approached. 


Discussion 

Arctic Greenland Shark populations have high 
infection rates with the ocular copepod parasite Om- 
matokoita elongata (Grant, 1827), which has been 
found in up to 98.9 percent of individuals in surveyed 
populations (Berland 1961), and in 100 percent of 
individuals in a recent tracking study off Baffin Island 
(Skomal and Benz 2004). Based on histopathological 
findings in copepod-parasitized eyes, Borucinska et 
al. (1998) postulated that sharks infected with ocular 
Ommatokoita were likely to be blind. However, the 
sharks we observed were free bilaterally of external 
signs and lesions of this parasite, had clear corneal 
epithelium, showed definite visual orientation to divers, 
tracked divers with horizontal and vertical eye move- 
ment, repeatedly demonstrated visual avoidance of 
divers and objects on the bottom, and repeatedly exhi- 
bited the described behaviour when closely approached. 
Vision is thought to be a major sense for investigative 
and social behaviour, prey recognition and predation 
motor patterns in the majority of shark species (Gruber 
1977). The visual acuity of the non-parasitized sharks 
we observed in the St. Lawrence River may have pro- 
found implications for the social behaviour and pat- 
terns of predation of this population. The reason these 
sharks have relatively low ocular copepod infestation 
rates compared to Arctic populations is unclear, and 
may relate to parasite abundance and ecology, host fac- 
tors such as population density, nutrition and immune 
status, and/or environmental conditions such as dis- 
persion currents, seawater salinity and temperature. 

Our perception of this species as a predator of benth- 
ic fishes and a sluggish, blind, olfactory scavenger of 
marine mammal carcasses, with an important ecological 
role as a mammal “carcass opening” species in Arctic 
deep water ecosystems (Snelgrove and Smith 2002) 
has changed as a result of these first-hand observations. 
Though descriptions of active predation by Greenland 
Sharks have not been reported in the scientific litera- 
ture, there is evidence that this species is capable of 
actively stalking and killing marine mammals as well 
as scavenging the carcasses of marine mammals. Ri- 
doux et al. (1998) documented a freshly killed juve- 
nile Ringed Seal, Phoca hispida (Schreber, 1775), with 
bite marks across the chest, found in the stomach of a 
4.5 m Greenland Shark off Iceland. Our underwater 


HARVEY-CLARK, GALLANT, AND HART: VISION IN ST. LAWRENCE RIVER GREENLAND SHARKS 


357 


FIGURE 4: Male Greenland Shark posturing, exhibiting curva- 
ture of the dorsum, having just shown rapid downward 
flexion of head and tail, contacting the silt-covered 
substrate, which caused the visible cloudiness ahead of 
and behind the animal. This motor pattern was elicited 
in response to presence and close proximity of divers. 


observations support the contention that large, stealthy 
Greenland Sharks are capable of rapid manoeuvring, 
moderate speed swimming and potential predation of 
infirm, sleeping (Ridgway et al. 1975), or predator- 
naive juvenile seals. This population of sharks is visual 
and can avoid or seek divers visually, and using other 
senses in conditions of poor visibility, and we advise 
caution when diving under low visibility conditions in 
areas where these sharks are known to occur. St. Law- 
rence River Greenland Sharks remain in the same shal- 
low water area for multiple days, repeatedly revisit 
novel potential prey items in the environment, ascend 
from the bottom to investigate divers in mid-water, 
circle and maintain constant visual contact with divers 
rather than fleeing, and exhibit unusual motor display 
patterns when approached closely by divers. These 
findings are consistent with the behaviour of a shark 
species which has the potential to act as an oppor- 
tunistic predator as well as a scavenger. 


Acknowledgments 
The authors gratefully acknowledge the assistance 
of Alain Simard, Jean-Yves Forest and Sylvain Sirois. 


Literature Cited 

Berland, B. 1961. Copepod Ommatokoita elongata (Grant) 
in the eyes of the Greenland Shark — a possible cause of 
mutual dependence. Nature 191: 829-830. 

Bigelow, H. B., and W. C. Schroeder. 1953. Fishes of the Gulf 
of Maine. Bulletin of the United States Fish and Wildlife 
Service 53(74): 53-55. 

Borucinska, J. D., G. W. Benz, and H. E. Whitely. 1998. 
Ocular lesions associated with attachment of the parasitic 
copepod Ommatokoita elongata (Grant) to corneas of 
Greenland sharks, Somniosus microcephalus (Bloch and 
Schneider). Journal of Fish Diseases 21: 415-422. 

Caloyianis, Nick. 1998. Greenland Sharks. National Geo- 
graphic 194(3): 60-72. 


358 


Compagno, L. V. J. 1984. Sharks of the World. An annotated 
catalogue of shark species known to date. FAO Fisheries 
Synopsis 125. Hexanchiformes to Lamniformes 4: 103-105. 

Gruber, S. H. 1977. The vision of sharks: a perspective. Naval 
Research Reviews 30(2): 1-27. 

Herendorff, C. E., and T. M. Berra. 1995. A Greenland 
Shark from the Wreck of the SS Central America at 2200 
Metres. Transactions of the American Fisheries Society 
124: 950-953. 

Lucas, Z. N., and D. F. McAlpine. 2002. Extralimital occur- 
rences of Ringed Seals, Phoca hispida, on Sable Island, 
Nova Scotia. Canadian Field-Naturalist 116: 607. 

Ridgway, S. H., R. J. Harrison, and P. L. Joyce. 1975. Sleep 
and cardiac rhythm in the grey seal. Science 187 (4176): 
933-359. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Ridoux,V., A. J. Hall, G. Steingrimsson, and G. Olaffson. 
1998. An inadvertent homing experiment with a young 
ringed seal, Phoca hispida. Marine Mammal Science 14: 
883-888. 

Skomal, G. B., and G. W. Benz. 2004. Ultrasonic tracking of 
Greenland sharks, Somniosus microcephalus, under Arctic 
ice. Marine Biology 145: 489-498. 

Snelgrove, P. V. R., and C. R. Smith. 2002. A riot of species 
in an environmental calm: the paradox of the species-rich 
deep-sea floor. Oceanography and Marine Biology Annual 
Review 40: 311-342. 


Received 6 November 2003 
Accepted 10 May 2005 


Charactéristiques d’une population introduit du Grand brochet, 
Esox lucius, dans le lac Ramsay, Parc de la Gatineau, Québec, 
et impact sur l’ichtyofaune 


JOSIANE VACHON!, BRIGITTE F. LAVALLEE, et FRANCOIS CHAPLEAU 


Département de biologie, Université d’Ottawa, Casier Postal 450, Succursale A, Ottawa, Ontario KIN 6N5 Canada 
' Auteur correspondant: jvach026 @alumni.uottawa.ca 


Vachon, Josiane, Brigitte F. Lavallée, et Francois Chapleau. 2005. Charactéristiques d’une population introduit du Grand 
brochet, Esox lucius, dans le lac Ramsay, Parc de la Gatineau, Québec, et impact sur l’ichtyofaune. Canadian Field- 
Naturalist 119(3): 359-366. 


En 1995, la présence du Grand brochet (Esox lucius), une espéce piscivore, a été notée pour la premiere fois dans le lac Ramsay, 
Parc de la Gatineau (Québec). Il a été déterminé que I’espéce a été introduite apres |’ été 1991. Un échantillonnage exhaustif de 
Vichtyofaune du lac en 2001 et 2002 a permis de constater que trois des 17 petites especes de poissons du lac sont probablement 
disparues: le Mulet perlé (Margariscus margarita), l’Epinoche a cing épines (Culaea inconstans) et une forme rare de l’Epinoche 
a trois épines (Gasterosteus aculeatus). Il est prédit que plusieurs autres espeéces pourraient disparaitre au cours des prochaines 
années. La croissance du Grand brochet dans le lac Ramsay est comparable aux autres lacs du sud du Québec. En 2001, le 
rapport femelles:males était de 15 : 1, une valeur anormale et inexpliquée. 


Mots clés: Esox lucius, Grand brochet, introduit, piscivore, croissance, petits lacs, Parc de la Gatineau. 


In 1995, the Northern Pike (Esox /ucius), a piscivorous fish, was captured for the first time in Ramsay Lake, a small lake of the 
Gatineau Park (Québec). It was determined that this species was introduced after the summer of 1991. An exhaustive survey 
of the ichthyofauna of this lake in 2001 and 2002 indicated that three of the 17 small-bodied species of the lake have proba- 
bly disappeared: the Pearl Dace (Margariscus margarita), the Fivespine Stickleback (Culaea inconstans) and a special form 
of the Threespine Stickleback (Gasterosteus aculeatus). It is predicted that several other species may be extirpated in the next 
few years. Northern Pike growth in the lake is comparable to other values obtained for other southern Québec lakes. In 


2001, the female:male ratio was highly unbalanced and difficult to explain at 15: 1. 


Key Words: Esox lucius, Northern Pike, introduction, piscivory, growth, small lakes, Gatineau Park. 


Lintroduction d’espéces exotiques de poissons per- 
turbe les communautés animales aquatiques (He et 
Kitchell 1990; Chapleau et al. 1997; Knapp et Mat- 
thews 2000; Economidis et al. 2000; Rahel 2000; 
Huckins et al. 2000; Findlay et al. 2000). Les petits 
lacs nordiques sans poissons piscivores de |’ Amérique 
du nord comptent beaucoup moins de petites especes 
de poissons que les lacs avec piscivores (Tonn et 
Magnuson 1982; Robinson et Tonn 1989; Tonn et al. 
1992). De plus, l’introduction d’une espéce piscivore 
dans un lac sans piscivore entraine inexorablement la 
disparition de plusieurs petites espeéces (Chapleau et al. 
1997; Whittier et al. 1997; Whittier et Kincaid 1999; 
Findlay et al. 2000). Ces espéces sont éliminées des 
petits lacs parce que toutes les classes d’ age sont vul- 
nérables a la prédation et qu’elles ne peuvent pas trou- 
ver un refuge contre |’action prédatrice des poissons 
piscivores (Tonn et al. 1992). Le cadre temporel du 
processus d’extinction locale suite a |’ introduction de 
piscivores n’a été que peu étudié (voir Demers et al. 
2001a and 2001b). En effet, la plupart des études n’ ont 
documenté que le fait accompli, souvent plusieurs 
décennies apres |’introduction initiale des piscivores 
(Chapleau et al. 1997; Whittier et Kincaid 1999; 
Findlay et al. 2000). 


Le lac Ramsay est un lac de 10,3 hectares qui a été 
inclus dans une étude sur |’impact de l’introduction 
(vers 1908) d’espéces piscivores sur les communautés 
de poissons des petits lacs du Parc de la Gatineau 
(Chapleau et al. 1997). En 1991, ce lac sans piscivores 
contenait 17 especes dont 12 espéces de petite taille 
(incluant 10 cyprinidés). De fait, ce lac présentait 
lichtyofaune la plus diversifiée des petits lacs (moins 
de 60 hectares) du Parc de la Gatineau. A l’automne 
1995, des spécimens d’une espéce piscivore, le Grand 
brochet (Esox lucius), ont été capturés dans le lac Ram- 
say. Le plus vieux spécimen étant agé de cing ans, nous 
constatons donc qu’une introduction non-autorisée de 
Grands brochets a été faite apres l’été 1991. Depuis, 
des échantillonnages (1998, 1999, 2001 et 2002) ont 
révélé que le Grand brochet était bien établi dans le 
lac Ramsay. 

Le premier objectif de notre étude est de déterminer 
si introduction d’une espéce piscivore apres |’ été 1991 
a eu un impact sur le nombre d’especes de poissons de 
petite taille dans le lac Ramsay. Un échantillonnage 
intensif des espeéces du lac en 2001 et 2002 a permis 
de répondre a cette question. Le deuxieme objectif de 
notre étude est d’examiner les paramétres biologiques 
de la population de Grand brochet (croissance, coeffi- 


o39 


360 


cient de condition, ratio des sexes) dans le lac Ramsay. 
Il est postulé que le Grand brochet montrera une crois- 
sance particuli¢rement rapide dans ce lac du fait qu’il se 
retrouve dans un habitat qui posséde une grande abon- 
dance de petits poissons et aucun autre piscivore. 


Matériel et méthode 

Le lac Ramsay (45°35'54"N, 76°06'00"0; profon- 
deur maximale : 9,5 m; profondeur moyenne : 4,1 m) 
est un petit lac du bouclier canadien a 200 m au-dessus 
du niveau de la mer. Son littoral est caractérisé par une 
abondance de plantes aquatiques mais également par 
des zones couvertes de sphaigne ainsi que des plages 
de sable et gravier. Le site est fréquenté irréguli¢rement 
par des campeurs |’été et des pécheurs |’été comme 
lV hiver (observation personnelle). 

Le tableau 1 énumere les 17 espéces de poissons 
trouvées dans le lac en 1991 (Chapleau et al. 1997). 
Cette liste provient d’un échantillonnage fait en 1991 
et est combinée a une liste d’especes publiée par Rubec 
(1975). En 1991, le lac contenait une forme relative- 
ment rare d’Epinoche A trois épines (Gasterosteus 
aculeatus). Il s’agit d'une forme habituellement ana- 
drome qui présente une série de plaques osseuses le 
long du corps et qui a pénétré dans la région en méme 
temps que la mer Champlain (McAllister et Coad 
1974). Elle est également restée prisonniére des lacs 
Hawley, Kidder, Pink et Canard suite au retrait des 
glaciers (Wisconsin) (Rubec 1975). 

Des échantillonnages exhaustifs de la communauté 
de poissons du lac Ramsay ont été faits en 1971 (Rubec, 
1975), 1991 (Chapleau et al. 1997), 2001 et 2002. En 
1995 et 1998, l’échantillonnage était surtout axé vers 
la capture de Grands brochets. En 1995, des Grands 
brochets furent capturés au filet maillant dans le cadre 
d’un cours de biologie (Université d’Ottawa). Les me- 
sures ainsi que les cleithra des poissons ont été con- 
servés. En 1998, les engins de péche comprenaient: 
trois filets maillants expérimentaux (50 m x 1,8 m 
avec mailles variant entre 2,5 et 15 cm) et un filet tré- 
mail (37 x 1.2 m; panneau extérieur avec mailles de 
20 cm et panneau intérieur avec mailles de 2,5 cm) et 
deux verveux (aile de 9,1 m x 0,9 m avec anneaux de 
76 cm de diamétre, mailles de 6 mm). En février 1999, 
20 brimballes ont été utilisées pendant deux jours 
(8 heures par jour). Elles étaient réparties également 
dans les deux sites ot les captures ont été les plus nom- 
breuses |’automne précédant. Un seul Grand brochet 
a été capturé lors de la péche hivernale. Ce spécimen 
n’a pas été inclus dans les analyses. En 2001, nous 
avons utilisé les mémes filets qu’en 1998 et la période 
de péche active a été de 72 heures (24 heures du 19 au 
21 septembre et 48 heures du 27 au 29 septembre). 
Tous les Grands brochets capturés ont été sacrifiés, 
rapportés au laboratoire dans des glaciéres et congelés. 
Les spécimens d’autres especes capturées dans les 
filets (Barbotte brune, Ameiurus nebulosus, et Meunier 
noir, Catostomus commersoni) ont été remis a |’ eau. 
L’échantillonnage des poissons de petite taille (surtout 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


des cyprinidés) en 2001 a été fait en utilisant une seine 
a batons de 10 m de longueur et deux verveux (dia- 
metre de l’ouverture: 76 cm; ailes 9 m). En 2002, seule- 
ment la seine a été utilisée. Les poissons de petite taille 
ont été remis a l’eau sauf pour quelques spécimens qui 
ont été conservés pour la confirmation d’ identification 
en laboratoire. Un seul Grand brochet (0+) a été cap- 
turé et il n’est pas inclus dans les analyses. L effort 
d’échantillonnage des petites especes en 2001 et 2002 
était plus élevé qu’en 1991. En 1991, nous avons échan- 
tillonné une surface de 300 m7? avec la seine alors que 
les verveux ont été actifs pendant 260,25 heures. En 
2001 et 2002, l’échantillonnage par la seine a couvert 
une surface de 3043 m/? alors que les verveux ont été 
laissés dans l’eau pendant 70,25 heures. II faut noter 
qu’aucune espece n’a été échantillonnée uniquement 
par les verveux en 1991, 2001 et 2002, ce qui suggére 
que la seine est le meilleur engin de péche pour la cap- 
ture des petites espéces. Nous avons donc échantillon- 
né une surface 10 fois plus grande en 2001 et 2002 
qu’en 1991. 

Tous les Grands brochets ont été décongelés avant 
d’étre pesés et mesurés (longueurs totale et standard). 
Un examen visuel des gonades a permis d’ identifier le 
sexe des spécimens. Le contenu stomacal a été exam- 
iné et les proies ont été identifiées lorsque cela était 
possible. Les deux cleithra ont été disséqués et bouil- 
lis. La détermination de |’ age a été faite en comptant 
les annuli sur les cleithra selon la méthode décrite par 
Casselman (1979). Les ages ont été confirmés par deux 
auteurs. Afin de faire des rétrocalculs de la longueur 
des individus a tous les ages, les distances entre le 
foyer et les annuli ainsi qu’entre le foyer et le bord des 
cleithra ont été mesurées. 

Le coefficient de condition des populations a été 
déterminé en calculant la pente de la relation entre les 
logarithmes de la masse et de la longueur totale (Tesh 
1971 et Ricker 1979 dans Wootton 1998). 


Résultats 
Ichtyofaune 

En 2001 et 2002, nous avons capturé 14 especes de 
poissons (sans compter le Grand brochet), soient trois 
de moins que lors de I’échantillonnage de la commu- 
nauté de 1991 alors qu’il n’y avait pas de Grands bro- 
chets dans le lac (tableau 1). Les espéces manquantes 
sont le Mulet perlé (Margariscus margarita), lEpi- 
noche a trois épines (Gasterosteus aculeatus) et \E- 
pinoche a cinq épines (Culaea inconstans). Certaines 
especes ont été capturées en trés petit nombre : le Méné 
laiton (Hybognathus hankinsoni), le Ventre citron 
(Phoxinus neogaeus) et V hybride entre le Ventre rouge 
du nord (Phoxinus eos) et le Ventre citron. Cependant, 
les difficultés d’identification des spécimens apparte- 
nant au complexe Phoxinus eos X Phoxinus neogaeus 
nous empéchent de conclure quoi que ce soit a propos 
de ces espéces. Les espéces les plus communes dans 
le lac sont la Barbotte brune (Ameiurus nebulosus), le 
Crapet-soleil (Lepomis gibbosus), la Chatte de lest 


2005 


VACHON, LAVALLEE, ET CHAPLEAU : GRAND BROCHET DANS LE LAC RAMSAY 361 


TABLEAU |. Liste des espéces de poissons dans le lac Ramsay, Parc de la Gatineau, en 1991 et en 2001-2002. 


Espéces capturées lors de I'échantillonnage de 199] 
Nom latin 


Grand brochet 
Umbre de vase 
Méné laiton 

Chatte de l'Est 


Esox lucius! 

Umbra limi 
Hybognathus hankinsoni 
Notemigonus crysoleucas 
Luxilus cornutus 


Notropis heterolepis Museau noir 


Phoxinus eos? Ventre rouge du nord 


Ventre citron 
Ventre-pourri 
Téte-de-boule 
Mulet a cornes 
Mulet perlé 
Meunier noir 
Barbotte brune 
Crapet-soleil 


Phoxinus neogaeus* 
Pimephales notatus 
Pimephales promelas 
Semotilus atromaculatus 
Margariscus margarita 
Catostomus commersoni 
Ameiurus nebulosus 
Lepomis gibbosus 
Etheostoma exile 
Culaea inconstans 
Gasterosteus aculeatus 


' Esox lucius n’a pas été capturé en 1991. 


Nom commun 


Méné a nageoires rouges 


Dard a ventre jaune 
Epinoche a cing épines 
Epinoche a trois épines 


Especes présentes 
en 2001 et 2002) 


a Eo oc oo 


mK OK 


2 S’hybride de Phoxinus eos et de Phoxinus neogaeus était également présent lors de cets échantillonnage. 


(Notemigonus chrysoleucas) et le Meunier noir (Cato- 
stomus commersoni). 

Parmi les Grands brochets capturés en 2001, 28% 
avaient l’estomac vide, 41% avaient un contenu digéré 
et 31% contenaient des poissons ou autres animaux 
non digérés. Trois des Grands brochets avaient des 
parties d’écrevisses, un avait capturé un petit rongeur et 
un autre une grenouille. Parmi les poissons consommés 
encore identifiables, on a noté la présence de Chattes 
de l’est et de Museaux noirs. Ces derni€res espéces 
ont été capturées lors de notre échantillonnage. 


Croissance du Grand brochet 

A l’automne 1995, 1998 et 2001, nous avons capturé 
respectivement 11, 34 et 32 Grands brochets. Le nom- 
bre de captures par unité d’ effort de péche (ou capture/ 
heure de péche active xX nombre de filets) de 1998 et 
2001 étaient respectivement de 0,095 et 0,099. Le nom- 
bre de Grands brochets capturés par unité d’effort (UE) 
a diminué lors des deux derniers jours de péche au filet 
maillant en 2001. Il était de 0,15 et 0,186 Grand brochet 
par UE pour les deux premiers jours de péche, puis de 
0,03 Grand brochet par UE lors des deux derniers jours. 

Tous les individus étaient agés entre 0+ et 7+ 
(Tableau 2). Les cleithra dont le premier annulus n’ était 
pas visible n’ont pas été utilisés pour le rétrocalcul de 
la longueur a l’age d’un an. Les pentes des régressions 
entre la longueur et la masse (coefficient de condition) 
étaient de 3,14, 3,07 et 2,93 pour 1995, 1998 et 2001 
respectivement (figure 1). Lors de la péche hivernale 
de 1999, le seul Grand brochet capturé était 4gé de deux 
ans. En juillet 2002, un Grand brochet 0+ (longueur 
totale = 12,4 cm, masse totale = 9,8 g) a été capturé 
avec la seine. 


Les longueurs moyennes rétrocalculées des Grands 
brochets a chaque age et a chaque année de capture 
sont présentées a la figure 2. Il y a généralement une 
différence significative entre les longueurs moyennes 
rétrocalculées des Grands brochets pour chaque année 
de capture, sauf lorsque les spécimens étaient agés de 
trois ans (ANOVA: alpha = 0,05; F = 0,528; p = 0,593). 
A un an, toutes les longueurs moyennes étaient signi- 
ficativement différentes (ANOVA: alpha = 0,05; F = 
60,105; p < 0,000). A deux ans, les Grands brochets 
capturés en 2001 avaient une longueur moyenne signi- 
ficativement plus élevée que les spécimens capturés 
en 1995 ou 1998 (ANOVA: alpha = 0,05; F = 12,912; 
P < 0,000: P ponterron de 1995 et 1998 < 0,000). A quatre 
et cing ans (ANOVA,,,,,: alpha = 0,05; F = 5,765; p = 
0,007 et ANOVA,,,.: alpha = 0,05; F = 8,058; p = 
0,003), la longueur moyenne des Grands brochets 
capturés en 2001 était significativement supérieure a 
celle des spécimens capturés en 1998 (ANOVA,,,.: 
alpha = 0,05; p ponferroni = 0,007; ANOVA.,,.: alpha = 
0,05; P Bonferroni = 0,002). 


Rapport femelles : males (F : M) 

En 2001, sur un total de 32 Grands brochets cap- 
turés, seulement deux étaient des males (rapport F:M 
de 15) 2,1). Ce\rapport, était de 1,2): lien 1995, alors 
qu'il était a de 5,8 : 1 en 1998. 


Discussion 
Ichtyodiversité 

Le Grand brochet a été introduit dans le lac Ramsay 
apres l’eté 1991. Depuis ce temps, trois especes de pois- 
sons (Mulet perlé, Epinoche a trois épines et Epinoche 
a cing épines) sont probablement disparues du lac 


362 THE CANADIAN FIELD-NATURALIST Vol. 119 


TABLEAU 2. Longueur et masse totales moyennes, minima et maxima pour chaque année d’échantillonnage et chaque classe 
d’age pour les Grands brochets (Esox lucius) du Pare de la Gatineau. Le nombre de captures (femelles : males) est égale- 
ment indiqué. 


Age Longueur Longueur totale Masse Masse totale 
(ans) totale min — moyenne + totale min — moyenne + Femelles 
Année max (cm) écart-type (cm) max (g) écart-type (g) : Males 
1995 3+ 52.0 — 67.7 56.7 =6.8 983.0 — 2419.0 1561.5 + 591.4 4:2 
4+ 50.0 — 72.0 61 fo 10:7 1006.0 — 2947.0 1918.0. 2952.3 232 
5+! - oly, ~ 1939.0 Od 
1998 I+ 33,5 > 31.1 35.4+ 1.8 230.0 — 309.0 Die ee ae 3.50 
2+ 34.7 — 41.2 36.1 = 22 271.0 — 444.0 353:8. 2 Det 10.52 
3+ 56.4 — 58.4 574+ 1.4 1179.0 — 1317.0 1248.0 + 97.6 20 
4+ 56.1 = 63.3 © pe ae a | 1222.0 — 1645.0 1368.3°2 239/77 Ae 
5+ 58.3 — 64.8 61.8: "2:2 1330.0 — 1879.0 1548 +\9922 Bz 
6+ 56.0 — 58.5 a/3 ELS 1092.0 — 1423.0 1257.9) 2340 220 
7+ 66.5 — 71.1 68.84 3.2 1890.0 — 2427.0 2158S te SIO. Zeal 
1939 3+! - 49.5 — 786 Od 
2001 1+! - 41.5 - 427.0 LO 
2+ 42.4 — 50.5 46.2+ 2.3 445.0 — 871.0 605.6 = 115.1 13.4.1 
3+ 51.0 — 54.0 52.2, , 1.3 807.0 — 1014.0 $53.0 + 973 P| 
4+ 58.5 — 68.9 64.5+ 4.5 873.0 — 2213.0 1502.0 + 546.9 5:9 
5+ 64.2 — 71.7 67.3.2. 2:8 1549.0 — 2308.0 1862.9 + 288.1 8:0 
2002 0+! - 12.4 - 9.8 1 n/d 


' Ces classes d’Age n’ont qu’un seul poisson. 


3.6 
1995: log masse = 3.14 log longueur -2.38 
1998: log masse = 3.07 log longueur - 2.30 
2001: log masse = 2.93 log longueur - 2.11 
Bee 
D 
© 
£ 
= 
A: 
2) 
Oo 
= 
0) 
O 
a 
2.4 
Z 


1.8 


1s Tf, ae 


Log longueur totale (cm) 


1.6 


FIGURE |. Relation logarithmique entre la masse totale et la longueur totale des Grands brochets du lac Ramsay. Le coefficient 
de condition correspond 4 la pente des droites. 1995 : n= 11 ; 1998 : n= 34; 2001 : n= 32. 


2005 


VACHON, LAVALLEE, ET CHAPLEAU : GRAND BROCHET DANS LE LAC RAMSAY 


363 


800 5 

600 + 
E 
Ss 
o 
wo 
3 
S 
° 
@ 400 4 
x) 
oO 
8 
5 
o 
& #1995 
9S #1998 

200 + 42001 

O + i= T T si] T Ur T 
1 2 3 4 5 6 7 8 


Age (ans) 


FiGurRE 2. Longueur rétrocalculée 4 chaque age des Grands brochets du lac Ramsay lors des années d’échantillonnage (1995, 
1998, 2001). Les barres d’ erreurs représentent les écarts-types. 


Ramsay. Leur présence dans le lac est possible mais 
dans ce cas, elle reste indétectable méme suite a un 
effort d’échantillonnage plus intensif en 2001 et 2002 
qu’en 1991. Il est probable que la cause de |’ absence 
de ces espéces soit |’ introduction de l’espéce piscivore. 
Le processus d’extinction des petites espéces est donc 
bel et bien enclenché dans le lac Ramsay et il semble 
rapide. Nous pouvons d’ores et déja prédire plusieurs 
autres extinctions au cours des prochaines années si la 
population de Grands brochets persiste. Chapleau et al. 
(1997) ont trouvé que les petits lacs du parc de la Gati- 
neau avec piscivores introduits ont moins de la moitié 
du nombre d’espéces de petite taille que les lacs qui 
n’ont pas de piscivores. Demers et al. (2001a) ont obser- 
vé que le nombre d’espéces non piscivores est passé de 
sept especes en 1991 a quatre en 1997 suite a |’ intro- 
duction, dans un petit lac, de deux espéces piscivores, 
l’Achigan a grande bouche (Micropterus salmoides) 
et l’Achigan a petite bouche (Micropterus dolomieu). 
Findlay et al. (2000) ont montré que le nombre d’es- 
peces de cyprinidés des lacs des Adirondacks était trois 
fois plus faible dans les lacs ayant des piscivores intro- 
duits. Nous pouvons donc prédire que la faune actuelle 
du lac Ramsay pourrait passer de 12 4 quatre ou six 
petites espéces au cours des prochaines années. 
D’ailleurs, les trois espéces disparues appartiennent 
a la liste des cing espéces présentes exclusivement 
dans les lacs sans espéces piscivores du Parc de la 
Gatineau (Chapleau et al. 1997). Les deux autres es- 
peces de cette liste, 1] Umbre de vase (Umbra limi), et 


le Méné laiton (Hybognatus hankinsoni) sont donc les 
prochaines candidates logiques a |’extinction locale. 
D’ailleurs, cette derniére espece n’a été capturée que 
trés rarement lors de notre échantillonnage alors qu’elle 
était abondante en 1991 (Chapleau, observation per- 
sonnelle). Le fait que les quatre espéces non pisci- 
vores les plus abondantes dans le lac en 2001 et 2002 
(Notemigonus chrysoleucas, Catostomus commersoni, 
Ameiurus nebulosus et Lepomis gibbosus) sont des 
espéces de taille intermédiaire démontre qu’il y a eu 
un changement de |’abondance relative des espéces, 
et cela en faveur des espéces de taille intermédiaire 
(Chapleau, observation personnelle). 

Demers et al. (2001a) ont démontré, suite a une 
manipulation des communautés de poissons dans deux 
petits lacs du sud de |’Ontario, que les stocks de pois- 
sons diminuaient fortement (de 32 kg ha! an! a5 kg 
ha! an‘') en trois a cing ans suite a l’introduction de 
piscivores. Dans un autre lac, ils ont également noté 
un certain rétablissement des stocks (de 12 kg ha’! 
an! 4 32 kg ha’! an’') en trois a cinq ans suite a |’en- 
lévement d’une partie des prédateurs. Dans ce méme 
lac, ils ont observé que deux especes saisonnicres sont 
devenues résidentes suite au retrait des Achigans a 
grande bouche et des Achigans a petite bouche. II serait 
donc possible de penser que le prélevement d’une par- 
tie importante de la population de Grands brochets en 
1998 et en 2001 ait ralenti le processus d’extinction et 
ait permis a certaines petites especes rares de se main- 
tenir. 


364 


De plus, en 1991, le lac Hawley, un petit lac (5,1 
hectares; 17 espéces; 12 espéces de petite taille) tout 
juste en aval (moins de | km) du lac Ramsay, ne comp- 
tait pas d’espéces piscivores. Lors d’un échantillonnage 
préliminaire en 1998, le Grand brochet a été signalé 
pour la premiere fois dans ce lac (Chapleau, observation 
personnelle). Donc, non seulement le Grand brochet 
est bien établi dans le lac Ramsay, mais il se répand 
présentement dans les lacs adjacents. 


Croissance du Grand brochet 

Les longueurs totales rétrocalculées des individus du 
lac Ramsay capturés en 1998 et 2001 sont comparables 
aux moyennes pour les populations du sud du Québec 
(Valliéres et Fortin 1988) (voir figure 3). Les Grands 
brochets capturés en 1995 ont eu une croissance lente 
lors de leur premiere année. 

Nous avions prévu une croissance rapide des Grands 
brochets au lac Ramsay a cause de |’absence d’ autres 
especes piscivores et de l’abondance de proies. Or, la 
croissance est normale. Cela peut étre attribuables a 
une compétition intraspécifique plus importante que 
prévue. En effet, la petite taille du lac a probablement 
contribué a l’atteinte d’une densité assez élevée de 
Grands brochets en peu de temps. Nos résultats ne 
concordent donc pas avec Stocek et al. (1999) qui ont 
observé une croissance tres rapide du Maskinongé 
(Esox masquinongy) suite a son introduction dans les 
eaux du Nouveau-Brunswick. Stocek et al. (1999) ont 
postulé que ce taux de croissance était da a la faible 
compétition et a l’abondance de proies. II est fort 
probable qu'une partie des Grands brochets capturés 
au lac Ramsay en 1995 soient nés dans un autre lac, 
ce qui expliquerait la faible longueur totale moyenne 
pour les individus a un et deux ans. 

Dans plusieurs cas, la longueur moyenne rétrocal- 
culée est significativement plus élevée chez les spéci- 
mens de Grands brochets capturés en 2001. On peut 
supposer que le nombre de grands spécimens a dimi- 
nué de facgon importante lors de la péche particuliére- 
ment intensive de 1998. Les survivants a cette péche 
qui ont été capturés en 2001 auraient eu une croissance 
particulierement importante. Cela peut s’expliquer en 
partie par une diminution de la compétition intraspé- 
cifique causée par la péche intensive. 

Les Grands brochets capturés en 2001 ont un coef- 
ficient de condition de 2,93 (Figure 2) ou une crois- 
sance allométrique négative, ils sont donc plus petits 
par rapport a leur longueur que la normale (c’est-a- 
dire, 3,0). De plus, leur condition semble a la baisse par 
rapport aux années d’échantillonnage précédentes. Les 
valeurs pour 1995 et 1998 étaient respectivement de 
3,14 et 3,07, soit une croissance allométrique positive, 
c’est-a-dire que les individus étaient généralement 
plus gros par rapport a leur longueur que la normale. 
Pour d’ autres populations, les valeurs obtenues se situ- 
ent habituellement entre 2,86 et 3,27 (Frost et Kipling 
1967; Mann 1976). [1 est difficile d’expliquer la dimi- 
nution du coefficient de condition dans le temps pour 


THE CANADIAN FIELD-NATURALIST 


Vol7aig 


les Grands brochets du Lac Ramsay. II est possible que 
l’abondance des proies montre une diminution dans le 
temps, la probable extinction de trois espéces en dix 
ans étant un bon indicateur que la présence du Grand 
brochet a eu un impact sur la communauté et possi- 
blement sur le recrutement des proies. 


Ratio femelles : mdles 

Le rapport femelles : males des Grands brochets 
évolue définitivement en faveur des femelles depuis 
1995. Tl est passé de 1,2 : len 1995; 45,8 : Dem 1998, 
pour finalement atteindre 15 : 1 en 2001. Ces derniéres 
valeurs sont tres élevées, mais au moins un autre résul- 
tat semblable a été observé. Frost et Kipling (1967) 
ont obtenu un ratio de 11,6: 1 avec les Grands brochets 
capturés a l’a4ge de quatre ans en 1951 au lac Winder- 
mere (données calculées a partir des résultats de Frost 
et Kipling 1967). Les années précédente et suivante 
montrent respectivement un ratio de 2,4: | et de 2,9: 1. 
Apres six ans, le ratio est revenu a | : 1. Les auteurs ne 
mentionnent pas ce débalancement. Dans la littéra- 
ture, les proportions varient le plus souvent entre 2 : 
1 et 1 : | (Frost et Kipling 1967 — calculé a partir des 
données de l’article; Casselman 1975; Mann 1976 — 
calculé a partir des données de I’article; Massé 1979 
dans Vallieres et Fortin 1988; Roche et al. 1999). 
Casselman (1975) a par ailleurs démontré que les pro- 
portions femelles: males capturés des Grands brochets 
avaient tendance a varier selon la saison. En été (crois- 
sance somatique) et en hiver (développement des go- 
nades), les femelles deviennent plus actives, ce qui les 
rend plus vulnérables a la prise au filet maillant. Or, 
chacun de nos échantillonnages a eu lieu vers la fin de 
l’été et au début de l’automne. La saison d’échantil- 
lonnage ne peut donc pas expliquer le ratio en faveur 
des femelles et l’évolution de ce ratio dans le temps. 
En septembre et octobre, les rapports femelles: males 
des populations étudiées par Casselman (1975) sont 
aux alentours de | : 1. Notre résultat suggére donc un 
sérieux débalancement dans la proportion des sexes 
dont la cause demeure inconnue. 


Remerciements 

Nous aimerions remercier le Parc de la Gatineau, 
la Commission de la Capitale Nationale et la Société 
de la Faune et des Parcs du Québec pour |’ attribution 
des permis de péche scientifique. Nous remercions 
également Anne-Marie Phelps, Thevia Belisle, Kevin 
Moon, Brent Campbell, Mélanie Barbeau, Annie Com- 
tois, Annie-Chantal Guibord, Julie Lepage et Guy 
Vachon pour leur aide avec |’échantillonnage. Ce pro- 
jet a pu étre complété grace a un octroi a la décou- 
verte du CRSNG a FC. 


Littérature Citée 

Casselman, J. M. 1975. Sex Ratios of Northern Pike, Esox 
lucius Linnaeus. Transactions of the American Fisheries 
Society |: 60-63. 


2005 


80 

PSB ONS 

= 

S 

co) 

© 

£ 49 : 
D | x 

= ab 

§ $ 


No 
Oo 
oK « 


VACHON, LAVALLEE, ET CHAPLEAU : GRAND BROCHET DANS LE LAC RAMSAY 


365 


ma< 


Lac Brochet 

* Sud du Québec 
@ Ramsay 2001 

mw Ramsay 1998 
@ Ramsay 1995 


FiGurE 3. Longueur rétrocalculée a chaque age des Grands brochets de lacs du Sud du Québec (Vallieres et Fortin 1988) 
incluant les données du lac Ramsay pour 1995, 1998 et 2001. 


Casselman, J. M. 1979. The esocid cleithrum as indicator 
calcified structure. Pages 249-272, édité par J. Dubé et Y. 
Gravel. Proceedings of the 10" Warmwater Workshop, 
Special Publications. Northeast Division, American Fish- 
eries Society. Montréal, Québec. 285 pages. 

Chapleau, F., S. Findlay, et E. Szenasy. 1997. Impact of 
piscivorous fish introductions on fish species richness of 
small lakes in Gatineau Park, Québec. Ecoscience 4: 259- 
268. 

Demers, E., D. J. McQueen, S. A. Popiel, A. M. Rocchi, C. 
W. Ramcharan, et A. Pérez-Fuentetaja. 2001a. Changes 
in fish community structure. Ergebnisse der Limnologie 
56: 23-48. 

Demers, E., D. J. McQueen, C. W. Ramcharan, et A. 
Pérez-Fuentetaja. 2001b. Did piscivores regulate changes 
in fish community structure? Ergebnisse der Limnologie 
56: 49-80. 

Economidis, P. S., E. Dimitriou, R. Pagoni, E. Michaloudi, 
et L. Natsis. 2000. Introduced and translocated fish species 
in the inland waters of Greece. Fisheries Management & 
Ecology 7: 239-250. 

Findlay, S. C., D. G. Bert, et L. Zheng. 2000. Effect of 
introduced piscivores on native minnow communities in 
Adirondack lakes. Canadian Journal of Fisheries and 
Aquatic Sciences 57: 570-580. 

Frost, W. E., et C. Kipling. 1967. A study of reproduction, 
early life, weight-length relationship and growth of pike, 
Esox lucius L., in Windermere. Journal of Animal Ecology 
36: 651-693. 

He, X., et J. F. Kitchell. 1990. Direct and indirect effects of 
predation on a fish community: a whole-lake experiment. 
Transactions of the American Fisheries Society 119: 825- 
835: 


Huckins, C. J. F., C. W. Osenberg, et G. G. Mittelbach. 
2000. Species introductions and their ecological conse- 
quences: An example with congeneric sunfish. Ecological 
Applications 10: 612-625. 

Knapp, R. A., et K. R. Matthews. 2000. Non-native fish in- 
troductions and the decline of the mountain yellow-legged 
frog from within protected areas. Conservation Biology 14: 
428-438. 

Mann, R. H. K. 1976. Observations on the age, growth, 
reproduction and food of the pike Esox lucius (L.) in two 
rivers in southern England. Journal of Fish Biology 8: 
179-197. 

McAllister, D. E., et B. W. Coad. 1974. Fishes of Canada’s 
National Capital Region. Fisheries Research Board of 
Canada Miscellaneous Special Publication 24. 200 pages. 

Rahel, F. J. 2000. Homogenization of fish faunas across the 
United States. Science (Washington, D.C.) 288 (5467): 
854-856. 

Robinson, C. L. K., et W. M. Tonn. 1989. Influence of envi- 
ronmental factors and piscivory in structuring fish assem- 
blages of small Alberta lakes. Canadian Journal of Fish- 
eries and Aquatic Sciences 46: 81-89. 

Roche, W., M. O’Grady, et J. J. Bracken. 1999. Some 
characteristics of a pike Esox lucius L. population in an 
Irish reservoir. Hydrobiologia 392: 217-223. 

Rubec, P. J. 1975. Fish distribution in Gatineau Park, Québec, 
in relation to postglacial dispersal, man’s influence and 
eutrophication. Canadian Field-Naturalist 89: 389-399. 

Stocek, R. F., P. J. Cronin, et P. D. Seymour. 1999. The mus- 
kellunge, Esox maskinongy, distribution and biology of a 
recent addition to the ichthyofauna of New Brunswick. 
Canadian Field-Naturalist 113: 230-234. 


366 


Tonn, W. M., et J. J. Magnuson. 1982. Patterns in the spe- 
cies composition and richness of fish assemblages in North- 
ern Wisconsin Lakes. Ecology 63: 1149-1166. 

Tonn, W. M., C. Paszkowski, et I. Holopainen. 1992. Pis- 
civory and recruitment: mechanisms structuring prey pop- 
ulations in small lakes. Ecology 73: 951-958. 

Vallieres, L., et R. Fortin. 1988. Le Grand brochet (Esox 
lucius) au Québec: biologie et gestion. Université du 
Québec a Montréal, pour le ministére du loisir, de la chasse 
et de la péche du Québec, Direction de la gestion des es- 
peces et des habitats, Québec. 298 pages. 

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Cyprinid distributions in Northeastern U.S.A. lakes: evi- 


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Vol. 119 


dence of regional-scale minnow biodiversity losses. Cana- 
dian Journal of Fisheries and Aquatic Sciences 54: 1593- 
1607. 

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Northeastern USA lakes: regional extent, dominance, 
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Series 24. London. 386 pages. 


Regu le 8 décembre 2003 
Accepté le 15 aott 2005 


Woodpecker Nest Tree Characteristics in Upper Midwestern 
Oak Forests 


COLLETTE L. ADKINS GIESE!? and FRANCESCA J. CUTHBERT 


Department of Fisheries, Wildlife, and Conservation Biology, 1980 Folwell Avenue, University of Minnesota, St. Paul, Min- 
nesota 55108 USA 

| Present address: 5415-134" Street North, Hugo, Minnesota 55038 USA 

? Corresponding author: e-mail: adki0020@umn.edu 


Adkins Giese, Collette L., and Francesca J. Cuthbert. 2005. Woodpecker nest tree characteristics in Upper Midwestern Oak 
forests. Canadian Field-Naturalist 119(3): 367-376. 


Characteristics of woodpecker nest trees have been widely studied in some regions of North America. However, there is little 
research from the Upper Midwest. Forest managers need information on woodpecker nest tree characteristics so they can recom- 
mend leaving during harvest trees that meet the needs of cavity-dwelling wildlife. Information specific to the Upper Midwest 
is especially important given that declines in several species of cavity nesting birds have been predicted by an environmental 
analysis of timber harvest in Minnesota. Our purpose was to identify attributes of nest trees used by primary cavity-nesting birds. 
We compared nest trees to unused trees and examined differences in nest trees among woodpecker species. We found 166 active 
woodpecker nests in upper midwestern oak forests in 1997 and 1998. For each nest tree, we recorded height, diameter, status, 
and aspects of tree decay. We also measured four potential nest trees (non-nest trees, within size requirements of cavity-nesting 
birds, with 22 indicators of heartwood decay) closest to each active nest tree. Additionally, we recorded these measurements for 
137 randomly selected potential nest trees. Using paired t-tests and chi-square analysis, we found each woodpecker species 
had a unique set of characteristics that separated nest trees from potential nest trees. Using an extension of the McNemar test 
for related samples, we found woodpeckers as a group used trees that were larger, both in diameter and height, more often 
elm (Ulmus americana, U. rubra) or aspen (Populus tremuloides, P. grandidentata), more likely to have old cavities present, and 
with more decay indicators than adjacent potential nest trees. The Yellow-bellied Sapsucker (Sphyrapicus varius) differed from 
the other woodpecker species by nesting in living Trembling Aspens (Populus tremuloides) with intact tops, complete bark 
cover, and heartwood fungus. Diameters of nest trees differed significantly among woodpecker species, but unlike findings 
from other studies, the height of nest hole and nest tree did not. Woodpecker nest entrances faced south or southeast signifi- 
cantly more often than by chance alone, even when excluding leaning trees. This study suggests that generic forest manage- 
ment for all woodpecker species may not be adequate because individual species have specific nest tree requirements. Man- 
agement recommendations for cavity-nesting birds need to be tailored to meet the needs of a diversity of species. 


Key Words: Woodpeckers, nest site selection, oak forests, timber management, Minnesota. 


Woodpeckers play a key role in forest communi- 
ties. The unique ability of woodpeckers to excavate 
holes in trees for nesting and roosting creates habitat 
for many other species of cavity-dwelling wildlife. 
Additionally, as predators of forest insects, woodpeck- 
ers may help control insect outbreaks (Bruns 1960). 
These natural history traits should make conservation 
of this group a priority in forest management. Declines 
in the numbers of several species [e.g., Barred Owl 
(Strix varia), Boreal Owl (Aegolius funereus), North- 
ern Saw-whet Owl (Aegolius acadicus), Yellow-bellied 
Sapsucker (Sphyrapicus varius), American Three-toed 
Woodpecker (Picoides dorsalis), Black-backed Wood- 
pecker (Picoides arcticus), Pileated Woodpecker (Dry- 
ocopus pileatus), Great-crested Flycatcher (Myiarchus 
crinitus), Black-capped Chickadee (Parus atricapillus), 
Boreal Chickadee (Poecile hudsonica), Redbreasted 
Nuthatch (Sitta canadensis), and White-breasted Nut- 
hatch (Sitta carolinensis)| of cavity-nesting birds have 
been predicted in the Upper Midwest (Generic Envi- 
ronmental Impact Statement on Timber Harvesting in 
Minnesota 1994, Minnesota Environmental Quality 
Board, St. Paul. 


Without knowledge of specific habitat requirements 
of cavity-nesting birds, conservation of these species 
is difficult. Information on woodpecker nest trees has 
important implications for forest management, includ- 
ing which trees are best to leave for wildlife during 
harvest and whether general guidelines are suitable for 
all species. 

Only certain trees are suitable for woodpecker ex- 
cavation. The tree must be large enough to support a 
nest, but selected tree size depends on woodpecker 
body size (Conner et al. 1975). In general, trees larger 
in diameter and height are used more often than small- 
er trees (Welsch and Howard 1983; Zarnowitz and 
Manuwal 1985). Woodpeckers also require trees with 
heartwood decay (Kilham 1971; Conner et al. 1976), 
which is caused by fungal invasion of the inner wood; 
decay softens the wood and facilitates excavation. 
Consequently, woodpeckers often choose dead or dying 
trees for nest hole excavation. However, some tree 
species (e.g., Populus spp.) can contain suitable heart- 
wood decay while they are still alive (Kilham 1971; 
Runde and Capen 1987). Woodpeckers are not restrict- 
ed to certain tree species for nesting. However, some 


367 


368 


" Chatfield Winona Co. 


Houston Co. 


MINNESOTA 


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THE CANADIAN FIELD-NATURALIST 


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WISCONSIN 


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FiGuRE |. Map of Houston and Fillmore counties of Minnesota and La Crosse County of Wisconsin showing locations of 
12 woodpecker nest tree study plots in upper midwestern oak forests, 1997-1998. 


tree species are preferred substrates for woodpecker 
nest excavation (Thomas et al. 1979). 

The characteristics of nest trees used by woodpeck- 
ers have been widely studied in some regions of North 
America. Research on woodpecker nest site selection 
in the western United States 1s most extensive (McClel- 
land and Frissel 1975; Bull and Meslow 1977; Scott 
1978; Scott et al. 1978; Mannan et al. 1980; Scott et 
al. 1980; Scott and Oldemeyer 1983; Winternitz and 
Cahn 1983; Raphael and White 1984; Zarnowitz and 
Manuwal 1985; Sedgewick and Knopf 1986, 1990; 
Li and Martin 1991; Schreiber and deCalesta 1992; 
Dobkin et al. 1995). 

Outside the western U.S., fewer researchers have 
examined characteristics of nest trees used by wood- 
peckers. Studies have been done in northern hard- 
wood forests of Vermont, New York, and New Hamp- 
shire (Kilham 1971; Swallow et al. 1986; Runde and 
Capen 1987). Other research was conducted in oak- 
hickory forests of southwestern Virginia (Conner 1975; 
Conner et al. 1975, 1976; Conner and Adkisson 1977). 
Only a few studies focused on the Upper Midwest, 
including oak-hickory forests of east-central Illinois 
(Reller 1972) and riparian areas in Iowa (Stauffer and 
Best 1982). 

Because characteristics of snags vary widely among 
biotic communities (Zeedyk 1983), information from 
other regions may not be applicable to upper mid- 
western oak forests. Geographically specific informa- 
tion on nest tree requirements of woodpeckers is need- 
ed to develop regional forest management guidelines 
for cavity-nesting birds. 

The objective of our study was to identify attributes 
of nest trees used by primary cavity-nesting birds 
(woodpeckers). We address the following questions: 
(1) Do trees chosen for nesting by woodpeckers differ 
from unused trees in terms of tree size, tree condition, 


and tree species? (2) Are there differences among 
woodpecker species in nest tree size, tree condition, 
and tree species? 


Methods 
Study Area 

Our study area was located in Houston (43°40'N, 
91°30'W) and Fillmore (43°40'N, 92°05'W) counties, 
southeastern Minnesota, and LaCrosse County 
(43°19'N, 91°27'W), western Wisconsin, United States 
(Figure 1). This area was uncovered by the Wiscon- 
sin Glaciation and is referred to as the Driftless Area. 
The pre-settlement vegetation was oak woodland and 
brushland and maple-basswood forest (Marschner 
1974). The landscape has become highly fragmented, 
consisting primarily of oak forest patches surrounded 
by agricultural lands. 

We selected plots from available state-owned forests 
(Richard J. Dorer Memorial State Hardwood Forest, 
Forestville State Park, and Coulee State Experimental 
Forest) on the basis of accessibility and intensity of 
public use. The plots were widely scattered: the west- 
ernmost and easternmost plots were approximately 
90 km apart. We studied 12 plots (8 plots each year; for 
maps see Adkins Giese 1999). Plots were 28-40 ha 
(mean = 36 ha). 

The study area has a varied disturbance history in- 
cluding logging, grazing, and fire. None of the plots 
was logged in the last ten years; however, firewood 
cutting was allowed at one site. Only the Quarry plot, 
which included some private land, was grazed in the 
last ten years. Some plots were in forest fragments as 
small as 40 ha, while others were within more exten- 
sive forest tracts. 

All plots were in closed-canopy mature forest, rang- 
ing in age from 80 to 120 years. The canopies were 
dominated by oaks (Quercus rubra, Q. alba, Q. bicol- 


2005 


or) (30 percent of all trees sampled), but also includ- 
ed hickories (Carya ovata, C. cordiformis) (13 per- 
cent), elms (17 percent), basswood (Tilia americana) 
(8 percent), aspen (6 percent), other hardwoods, and 
some White Pine (Pinus strobus). Young plants of these 
canopy trees and hazel (Corylus spp.), gooseberry 
(Ribes spp.), and raspberry (Rubus spp.) made up the 
woody understory. We provide additional details on 
habitat structure and composition in another paper 
(Adkins Giese and Cuthbert 2003), which focuses on 
the broader habitat needs of woodpeckers. Breeding 
bird surveys found that the plots had 56 bird species, 
including 13 cavity-nesting species (M. Friberg, Uni- 
versity of Minnesota, personal communication). 

We marked the plots with flagging tape, forming a 
grid used to plot locations of woodpecker activity on 
field maps, mark nest locations, and specify locations 
for the randomly selected sites. 


Nest Searching 

In 1997, we began searching for nests 6 May and 
stopped 23 June. In 1998, we began searching for nests 
20 April and stopped 22 June. After formal nest search- 
ing ended, we opportunistically located additional nests 
during vegetation surveys. 

We searched for active nests of all primary cavity- 
nesting birds on the plots, excluding the Black-capped 
Chickadee. The primary cavity-nesting birds included 
Downy (Picoides pubescens), Hairy (P. villosus), Red- 
bellied (Melanerpes carolinus), Red-headed (M. ery- 
throcephalus), and Pileated woodpeckers, Yellow- 
bellied Sapsuckers, and Northern Flickers (Colaptes 
auratus). 

We found nests by following woodpecker vocaliza- 
tions, drumming, and flight paths. We found addition- 
al nests by systematically walking the plots and exam- 
ining trees with cavities and fresh chips at the base. 
Our goal was to find as many nests as possible, rather 
than to make plot comparisons, thus there was no at- 
tempt to equalize nest-searching efforts across plots. 
We included nests found outside plots as long as they 
were within the study area. We found nests of the 
Red-headed Woodpecker along roadsides, as well as 
on the plots. 

We considered nests to be active if we observed 
any of the following: (1) adult completely entering 
cavity and remaining for over 10 minutes; (2) adult 
flushed from cavity; (3) adult feeding young; and (4) 
young calling from cavity. Because of difficulty locat- 
ing Red-headed Woodpecker nests, an adult repeatedly 
entering a cavity but not remaining inside was con- 
sidered sufficient evidence of nest activity. 


Vegetation Surveys 

We recorded characteristics of all active woodpeck- 
er nest trees found. For comparison, we also obtained 
vegetation measurements for the four closest potential 
nest trees and a sample of randomly selected potential 
nest trees. Potential nest trees were defined as non-nest 
trees, within the height and diameter requirements of 


ADKINS GIESE and CUTHBERT: WOODPECKER NEST TREE CHARACTERISTICS 


369 


cavity-nesting birds, with 22 indicators of heartwood 
decay (Conner 1978). Decay indicators included old 
cavities, tree scars, branch stubs, fungal conks, and 
significant dead portions. Potential nest trees were not 
presently used by woodpeckers, but some had indica- 
tions of previous use. In accordance with the minimum 
nest height and tree diameter used by woodpeckers, 
potential nest trees were >15.2 cm diameter at breast 
height (dbh) and >1.8 m tall (Thomas et al. 1979). We 
modified the study methods after Runde and Capen 
(1987). 

We recorded vegetation characteristics to describe 
tree size, tree condition, and tree species of nest trees 
and potential nest trees (Table 1). To describe tree size, 
we recorded tree height and dbh. To describe the tree 
condition, we recorded tree status, top condition, limb 
condition, presence of decay indicators, and percent- 
age live wood and bark cover in quartile classes. For 
consistency, the same person took all subjective veg- 
etation measurements. 

To determine the availability of trees of various sizes 
and species for nesting and to get a sample of random- 
ly selected potential nest trees, we sampled trees in 
circular subplots (11.3 m radius) randomly distributed 
across the plots. For all trees within the subplots, we 
recorded species, status, and size class. From subplots 
that contained potential nest trees, one potential nest 
tree was randomly chosen for comparison to nest trees 
Gr = 137). 


Data Analysis 

We compared nest trees to potential nest trees to 
evaluate woodpecker nest tree selection. Comparisons 
of nest trees among the woodpecker species were also 
made to determine interspecific similarities and differ- 
ences. Alpha levels < 0.05 were used to indicate sig- 
nificance. 

We statistically compared size, condition, and species 
of nest trees to adjacent potential nest trees. By using 
adjacent potential nest trees, rather than randomly 
selected potential nest trees, we could assess how the 
nest tree differed from other trees within the chosen 
nest area. We used paired f-tests to compare dbh and 
height of nest trees to the mean of four adjacent 
potential nest trees. Because data for available trees 
were categorical, with diameter based on size class, we 
used chi-square to compare the diameter of nest trees 
to available trees. An extension of the McNemar test 
for related samples (Miettinen 1968) was used to com- 
pare the condition and species of all woodpecker nest 
trees to the related sample of adjacent potential nest 
trees. Using the McNemar extension, we established 
comparisons using 2 or 3 categories for each independ- 
ent variable. 

Because of high skew and small sample size, the 
extension of the McNemar test could not be used when 
comparing the condition and species of nest trees to 
potential nest trees for each individual woodpecker 
species. Instead, we used chi-square tests of homo- 


370 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |. Descriptions of vegetation measurements taken for woodpecker nest trees and potential nest trees in upper mid- 


western oak forests, 1997-1998. 
Variable Categories 


Tree status Alive, Dead, Partly Dead 


Tree height N/A 
Top condition Top intact, Top broken, 
Broken fork 


Limb condition Trunk, Main branches, Secondary 


branches, Foliage twigs 


Percentage 0-25, >25-50, 
live wood >50-75, >75.. 
Percentage 0-25, >25-50, 
bark cover >50-75, >75 
Heartwood Presence, Absence 


fungal conks 
Sapwood fungus 
Old cavities 


Presence, Absence 
Presence, Absence 


Trees scars Presence, Absence 


Significant Presence, Absence 
dead portions 


Branch stubs Presence, Absence 


geneity to compare nest trees to a randomly selected 
sample of potential nest trees. Because each independ- 
ent variable had 2 or 3 categories, we set up 2 x 2 or 
2 x 3 chi-square contingency tables, accordingly. If 
we found a significant difference within a 2 x 3 chi- 
square contingency table, we broke the table into non- 
independent 2 x 2 tables for examination of where 
non-homogeneity occurred (Brunden 1972). 

We also made comparisons among the woodpeck- 
er species for nest tree size, condition, and species. 
We used Kruskal-Wallis ANOVAs to compare the 
dbh and height of nest trees among the seven species 
of woodpeckers. When we found a significant differ- 
ence among species, we used a multiple comparison 
procedure using rank sums to determine which species 
differed significantly from each other (Dunn 1964). 
To compare the condition and species of the nest tree 
among the species of woodpeckers, we used chi-square 
tests of homogeneity. The Northern Flicker and the 
Pileated Woodpecker were not included in compari- 
sons among the woodpecker species because of small 
sample size. 


Results 
Comparison of Nest Trees and Unused Trees 

We found 166 nests, including 76 in 1997 and 90 in 
1998 (Table 2). Woodpecker nest trees had significant- 
ly greater diameter and height than adjacent potential 
nest trees (Table 3). Over 50% of nest trees were 23- 
38 cm dbh (Figure 2). This diameter class was used out 
of proportion to its availability (y7 = 893, P < 0.001, 
Bl) 

The decay condition of active woodpecker nest trees 
also differed from potential nest trees. In comparison 


Description 


Partly dead: tree forked with only | fork dead or tree 
with only small remaining living branches (>75% dead) 

Measured using a clinometer 

Top broken: tree top ends abruptly. Broken fork: trunk 
split with | fork intact and | broken 

Smallest type of branches remaining, regardless of 
whether branches were dead or alive 

A subjective estimate of percentage live wood in 
quartile classes 

A subjective estimate of percentage bark cover in 
quartile classes 

Formes igniarius, the shelf fungus of Trembling Aspen, 
and other large shelf fungi 

Small, superficial tree fungi 

Holes that looked as if they were completely excavated 
by a woodpecker 

Tree wounds with exposed heartwood (natural cavities, 
excavation attempts, and deep foraging holes) 

Dead portions large enough to be nesting substrate for 
Downy Woodpecker (>15 cm diameter; >30 cm long) 
Broken branch or stem >15 cm diameter and >30 cm long 

or broken branch <50 cm on stem >15 cm diameter 


to adjacent potential nest trees, nest trees of all wood- 
pecker species combined were less likely to have brok- 
en tops (P < 0.05), were more likely to have foliage- 
bearing twigs present (P < 0.01), were more likely to 
have heartwood fungus present (P < 0.01), were less 
likely to have significant dead portions present (P<0.05), 
had more total decay indicators present (P < 0.001), 
and were more often elm or quaking aspen (P < 0.001). 
Nest trees also were more likely to have old cavities 
present (55% of nest trees versus 4% of potential nest 
trees), but the high skew invalidated the McNemar 
extension. These results were highly influenced by 
the large sample of Yellow-bellied Sapsuckers, which 
reflected the species’ strong preference for living aspens 
with intact tops and heartwood decay (Table 4). When 
the Yellow-bellied Sapsucker was excluded, only num- 
ber of decay indicators (P < 0.01) and tree species elm 
or aspen (P < 0.001) remained significant. The per- 
centage of nest trees with old cavities still remained 
high (51% of nest trees versus 8% of potential nest 
trees). There was no difference found between nest 
trees and potential nest trees in tree status, percentage 
live wood or bark cover, or in the presence of branch 
stubs or tree scars. 

Certain tree species were frequently used for nesting 
by woodpeckers. Woodpeckers nested in American Elms 
(Ulmus americana) and Trembling (Quaking) Aspens 
much more often than expected based on availability 
(7 = 391, P < 0.001, df = 1). American Elm and Trem- 
bling Aspen provided 70% of all nest sites, but con- 
stituted only 10% and 5% of all trees in random plots, 
respectively. Dead American Elms made up <1% of 
available trees and dead Slippery Elms (U/mus rubra) 
made up approximately 1% of available trees. 


a7] 


WOODPECKER NEST TREE CHARACTERISTICS 


ADKINS GIESE and CUTHBERT 


2005 


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S72, 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE 3. Paired comparison of nest trees with mean of four adjacent potential nest trees for seven species of woodpecker in 


upper midwestern oak forests, 1997-1998. 


Tree Diameter (cm) 


Ditf.* “SE t df 
Hairy Woodpecker -0.8 Zo SO 2h 
Downy Woodpecker -0.7 24 -0.8 43 
Yellow-bellied Sapsucker 0.1 fe Oa) ak 
Red-bellied Woodpecker 10.2 oe: TaN a ee 
Red-headed Woodpecker 17.0 ee > a 
Northern Flicker 314 ~ *20%6 1.6 3 
Pileated Woodpecker Bo OT a4 3 
All woodpecker nests sys) 6). 2:3. 164 


Tree Height (m) 
Diff. SE t df 

NS 2.8 13 2h 21 
NS? 2.6 | ol 2.4 43 ‘ 
NS 3.2 0.8 6.2 4] ay 
* 4.3 1.4 S| 28 = 
ee a 6.5 3 SH 19 ee 
NS 4.7 1.4 3.4 3 
* 9.4 6.2 3 s) NS 
lars 4.3 0.5 8.2 164 ie 


“ The difference of the means (Diff.) of nest tree and four adjacent potential nest trees (nest tree — adjacent) 


>NS P>0.05 
P2005 
“P<0.01 
P<) Oot 


100 


80 


m Nest Trees 
OD Available Trees 


60 
40 


20 


Percent of Trees 


8-15 
>15-23 Ff 
>23-38 F 
>38-53 | 
>53-69 
>69-84 
84-102 

>102 


Diameter at Breast Height (cm) 


FiGuRE 2. Diameter at breast height (cm) in size classes for 
all woodpecker nest trees and available trees in upper 
midwestern oak forests, 1997-1998. 


Each woodpecker species had a set of characteris- 
tics that differentiated its nest trees from randomly 
selected potential nest trees (Table 4). In comparison 
with random potential nest trees, nest trees of the 
Pileated Woodpecker were more often elm or Trem- 
bling Aspen (P < 0.001), with old cavities (P < 
0.001), more decay indicators (P < 0.001), and less 
bark cover (P < 0.01). Nest trees of the Northern 
Flicker were more often elm or Trembling Aspen (P 
< 0.01), with old cavities (P < 0.001) and more decay 
indicators (P < 0.001). 


Comparison Among Woodpecker Species 

We wanted to determine if nest tree selection dif- 
fered among species of woodpeckers. We found a 
significant difference among woodpecker species in 
nest tree dbh, but not nest tree height (Table 5). Many 
differences in nest tree condition among the wood- 
pecker species were found (Table 6). The Yellow-bel- 
lied Sapsucker differed from the other woodpecker 
species by nesting in living Trembling Aspens with 
intact tops, complete bark cover, and heartwood fun- 


gus. No difference among the woodpecker species 
was found in limb condition, number of decay indi- 
cators, and presence of branch stubs, tree scars, and 
old cavities. 


Discussion 
Tree Size 

Many investigators report that woodpecker nest 
trees are larger than unused snags (Bull and Meslow 
1977; Welsch and Howard 1983; Zarnowitz and Man- 
uwal 1985; Schreiber and deCalesta 1992). We com- 
pared nest trees to unused trees that met minimum size 
requirements. This allowed us to make a comparison 
with trees that are generally larger in diameter than a 
random sample of unused snags. We nevertheless found 
that woodpeckers chose trees that were larger in diame- 
ter than adjacent potential nest trees. There are many 
possible advantages to nesting in larger trees. Larger 
trees may contain more places to excavate. Larger trees 
are probably older and therefore more decayed. Addi- 
tionally, larger trees enable construction of cavities 
with thicker walls, which provide thermal insulation, 
protection from predators, and lower probability of 
breaking at cavity height (Kilham 1971; Miller and 
Miller 1980). 

Ranges of tree diameters used by Downy and Hairy 
woodpeckers and Yellow-bellied Sapsuckers in our 
study are comparable to ranges found in the literature 
(Conner et al. 1975; Evans and Conner 1979; Thomas 
et al. 1979). However, in our study, 55% of trees used 
by Red-bellied Woodpeckers and 20% of trees used by 
Red-headed Woodpeckers were below the observed 
optimum range of nest tree diameters given for these 
species (Evans and Conner 1979). Selection may be 
different for these species in our study area or our sam- 
ple may not be representative. However, it is possible 
that our results suggest a shortage in the study area of 
large diameter trees required by Red-bellied and Red- 
headed woodpeckers. 


2005 


TABLE 4. Characteristics that significantly differed (chi-square 
tests of homogeniety) for seven species of woodpecker in 
upper midwestern oak forests. 1997-1998. 


% of trees with 


characteristic 
Random 
Nest Potential 
trees Nest Tree* 
Downy Woodpecker (n = 44) 
Dead or partly dead’ 84 66 
Broken forked top” (d.f. = 2) 25 6 
Tree elm or Trembling Aspen’ 59 18 
Old cavities present™”* 4] 4 
Sapwood fungus present’ 32 1] 
3 or more decay indicators” 61 28 
0-25% bark cover’ 39 9 
Hairy Woodpecker (n = 22) 
Tree elm or Trembling Aspen’ 64 18 
Old cavities present 45 4 
Heartwood decay fungus present” 45 13 
Without significant dead portion” 36 9 
3 or more decay indicators” 68 28 
Red-bellied Woodpecker (n = 29) 
Tree elm or Trembling Aspen’ 52 18 
Broken forked top” (d.f. =2) 28 6 
Old cavities present’ 5) 4 
3 or more decay indicators” 66 28 
0-25% bark cover™ 31 9 
Red-headed Woodpecker (n = 20) 
Tree elm or Trembling Aspen’ ” 80 18 
Dead or partly dead” 90 66 
Broken fork’” (d.f. = 2) 30 6 
Old cavities present” 70 4 
3 or more decay indicators’ ~ 70 28 
0-25% bark cover’ 70 9 
Yellow-bellied Sapsucker (7 = 42) 
Tree alive" 88 34 
Top intact™™* (d.f. = 2) 98 62 
Tree Trembling Aspen’ ~ 88 8 
Old cavities present’”” 67 4 
Heartwood decay fungus present’ 95 13 
No significant dead portion’ ~~ 86 9 
3 or more decay indicators" 74 28 
75-100% bark cover* 95 81 


“n= 137 random potential nest trees 
> df. = 1 unless otherwise indicated 
~ P*=0.05 

we <0 

“P< 0.001 


Generally, woodpeckers with larger body sizes re- 
quire larger diameter nest trees (Conner et al. 1975). 
Similar to other studies (Conner et al. 1975; Brawn et 
al. 1984; Raphael and White 1984; Li and Martin 
1991), we found a significant difference among wood- 
pecker species in nest tree diameter, which generally 
corresponded with woodpecker body size. 

Woodpeckers chose nest trees that were taller than 
adjacent potential nest trees. When taller trees are avail- 
able, nest heights tend to be higher because higher nests 


ADKINS GIESE and CUTHBERT: WOODPECKER NEST TREE CHARACTERISTICS 


make nest cavities less easily detected and reached 
by predators (Miller and Miller 1980). A nest located 
high in the tree gives the woodpecker more time to 
dislodge or discourage a predator climbing the trunk 
(Kilham 1971). Other investigators have also found that 
woodpeckers choose taller trees (Welsch and Howard 
1983; Zarnowitz and Manuwal 1985; Sedgewick and 
Knopf 1990; Joy 2000). 

There were no significant differences among wood- 
pecker species in nest tree height or nest hole height 
in our study. However, other investigators have found 
differences in nest tree height or nest hole height 
(Conner et al. 1975; Stauffer and Best 1982; Brawn 
et al. 1984; Raphael and White 1984). Harestad and 
Keisker (1989) explained difference in nest tree height 
among woodpeckers as a consequence of different 
preferences for tree condition. 


Tree Condition 

Trees used by woodpeckers had several indicators 
of heartwood decay. Soft heartwood is a necessity for 
excavation (Kilham 1971), and tree wounds, such as 
branch stubs, tree scars, and old cavities, provide ac- 
cess to the heartwood and serve as entry points for 
fungi. Number of decay indicators present depended 
in part on the tree species. It appeared that tree species 
with especially hard wood, like oaks and cherries, 
required more decay for the wood to become suitably 
softened for excavation. 

Woodpeckers often chose trees with old cavities. 
Old cavities are clear indicators of past suitability 
and also serve as entry points for additional heart- 
wood fungi. Return of birds to the same tree is likely 
also a function of tree availability and territoriality. 
One active Yellow-bellied Sapsucker nest tree con- 
tained 16 old cavities. 

Differences in condition of trees chosen for nesting 
by each species may be explained by differences in 
the excavation abilities of the species (Spring 1965). 
Downy Woodpeckers are weak excavators and often 
choose trees with some sapwood fungi (Harestad and 
Keisker 1989; this study). Presence of sapwood fungi 
suggests that the outer wood is soft, making the wood 
easier to excavate. However, we found trees with exten- 
sive sapwood decay were not used for nesting. Such 
trees may not offer adequate protection of the nest 
cavity (Kilham 1971; Conner et al. 1976). Yellow-bel- 
lied Sapsuckers most often nested in living trees with 
sound sapwood and several indicators of heartwood 
decay. Fomes igniarius, a heartwood fungus that attacks 
live wood, especially of Trembling Aspen, softens the 
heartwood but leaves the sapwood sound (Harestad 
and Keisker 1989). Nearly all Yellow-bellied Sapsucker 
nest trees had conks of Fomes igniarius present. 


Tree Species 

American Elms and Trembling Aspens were most 
frequently used for nesting. Although we found no 
nests in Slippery Elms, there were more dead Slip- 


374 THE CANADIAN FIELD-NATURALIST Vol. 119 


TABLE 5. Diameter at breast height (dbh) and height of nest trees of seven species of woodpecker upper in upper midwestern 
oak forests, 1997-1998. 


Nest Tree dbh (cm) Nest Tree Height (m) 

n Mean SE Min Max Mean SE Min Max 
Yellow-bellied Sapsucker 42 30A 080 19 42 20A 0.62 8 34 
Hairy Woodpecker 22 33 A 3.0 20 76 I7A it 8 24 
Downy Woodpecker 44 35 AB* i 16 63 17A 0.91 6 29 
Pileated Woodpecker 4 69 BC 23 42 127 21A 50 9 29 
Northern Flicker 4 69 BC 22 38 121 I8A 2.6 14 23 
Red-bellied Woodpecker 20 47C 5.0 pe 137 18 A ot 8 20 
Red-headed Woodpecker 20 54C 7.0 28 168 19 A 2 Zi 545 
All woodpecker nests 101 I BAB: 16 168 18 0.5 6 34 


“Means with the same letter were not significantly different (P > 0.05) by multiple comparison procedure based on ranks 
TABLE 6. Significant differences (chi-square tests) among nest tree characteristics of five species of woodpecker in upper 
upper midwestern oak forests 1997-1998. 


% of nest trees with each characteristic 
Species Heartwood Sapwood Significant 


Trembling fungus fungus dead Livewood Bark cover Tree Tree Top 
n Aspen present present portion >75-100% >75-100% alive intact 
Downy Woodpecker 44 11 A? 16A 32 A 95A I8A 52 A 16A 48 A 
Red-bellied Woodpecker 29 11A 24 AC 28 AB 100 A 31A 66 A 24A 24A 
Red-headed Woodpecker 20 27A 10 AC 5 AB 90 AB 1OA 30A 1OA SOA 
Hairy Woodpecker 22 I7A 45C 14 AB 64 B 41 A 64A 36A 35 A 
Yellow-bellied Sapsucker 42 88&B 95 B 2B 14C 88 B 55 88 B 98 B 


“ Percentages with same letter reflect means that were not significantly different (P > 0.05) by multiple comparison. 


pery Elms available than dead American Elms. This 
observation suggests that American Elms have decay 
characteristics that suit woodpeckers. 

The propensity for Yellow-bellied Sapsuckers to nest 
in aspens has been well documented (Kilham 1971; 
Thomas et al. 1979; Runde and Capen 1987; Harestad 
and Keisker 1989). Aspens may be more prone to 
decay than most other hardwoods. In our study area, 
it appears that Yellow-bellied Sapsuckers are weak ex- 
cavators, as suggested by Jackman (1974), so the exten- 
sive heartwood decay of mature aspen are suitable. 

Red-headed Woodpeckers primarily nested in Amer- 
ican Elms, especially large, barkless elms with many 
broken branches. Dutch elm disease may have bene- 
fited Red-headed Woodpeckers by creating highly 
suitable snags (Jackson 1976). However, the disease 
widely eliminated American Elm as a major forest 
species, and in the future, woodpeckers will have few 
dead elms for nesting. Elms that died from Dutch 
elm disease are losing suitability for nesting as tree 
decay becomes extensive; additional dead elms are 
lost from blow-down and human removal. Red-head- 
ed Woodpeckers will need to rely more heavily on 
other tree species for nesting or face continued decline. 
Abundance of Red-headed Woodpeckers in Minnesota 
has been significantly declining since 1966 (Green 
1995): 


Downy, Hairy, and Red-bellied Woodpeckers showed 
more variation in tree species chosen. However, cau- 
tion should be observed before applying these tree 
species results to other geographic areas because tree 
species selected for nesting by these species vary by 
locality, availability, and tree condition (Bull et al. 
1980). 


Management Implications 

It is important that suitable cavity trees are left dur- 
ing timber harvest. The value of such trees extends 
beyond their importance to woodpeckers. In upper 
midwestern oak forests, secondary cavity-dwelling 
wildlife like the Great-crested Flycatcher, Tufted Tit- 
mouse (Baeolophus bicolor), House Wren (Troglo- 
dytes aedon), White-breasted Nuthatch, Barred Owl, 
and Southern Flying Squirrel (Glaucomys volans) all 
use old woodpecker holes for nesting. Studies have 
shown that lack of suitable nest sites is a limiting fac- 
tor for many species of cavity-nesting birds (Cunning- 
ham et al. 1980; Zarnowitz and Manuwal 1985; Dob- 
kin et al. 1995). 

Our study does not address number of trees to leave 
for wildlife during timber harvest, but it does offer 
insight into the characteristics of trees. According to 
our results, emphasis should be placed on retaining 
trees with sound sapwood that also show signs of 
heartwood decay (e.g. broken tops). We recommend 


2005 


retaining living as well as dead trees, but living trees 
need to be maintained on long rotations to develop 
suitable decay characteristics for nesting. Indeed, old 
growth conditions may take >100 years to develop in 
both oak-hickory forests and aspen forests (Hardin and 
Evans 1977; Winternitz and Cahn 1983). Because trees 
with old woodpecker nest cavities are especially im- 
portant for nesting, trees with old cavities should be 
retained. 

Trees of various size classes should be retained 
during harvest. It is important that diameters near the 
mean for each woodpecker species are provided to 
encourage excavation of normal-sized cavities and 
reduce death of nestlings due to overcrowding (Kil- 
ham 1968; Evans and Connor 1979; Miller and Miller 
1980). Small diameter trees may provide foraging sub- 
strate, but they should not be considered substitutes 
for larger diameter cavity trees. If choices need to be 
made about which trees to retain, the larger diameter 
and taller trees should be retained. 

In our study, certain tree species, especially Amer- 
ican Elm and Trembling Aspen, were used more often 
than others for nesting. However, undue emphasis 
should not be placed on importance of tree species. 
Most woodpeckers are restricted to certain tree species 
for nesting but require certain levels of decay, which 
vary by tree species. 

Our study suggests that generic management for 
all woodpecker species may not be adequate because 
individual species have specific nest tree requirements. 
However, it may be possible to meet a diversity of 
species needs if forest managers retain many large 
trees with a range of decay conditions. Although this 
paper focuses on the suitability of individual trees for 
woodpecker nesting, we discuss the role of the larger 
forest context in another paper (Adkins Giese and 
Cuthbert 2003). More research is needed on longevity 
of trees left for wildlife during harvest and the long- 
term impacts of existing forest management practices 
on wildlife. However, recommendations from this 
study, if adopted, would enhance conditions for cavi- 
ty-dwelling wildlife in upper midwestern oak forests. 


Acknowledgments 

The Minnesota Nongame Wildlife Tax Checkoff 
and the Reinvest In Minnesota Program, through the 
Minnesota Department of Natural Resources, Divi- 
sion of Fish and Wildlife, Natural Heritage and Non- 
game Research Program, provided funding for the 
fieldwork. Additional support came from Minnesota 
Forest Resources Council, the Gordon Gullion Schol- 
arship, and Dayton-Wilkie Natural History Fund of 
Bell Museum of Natural History, Minnesota Agricul- 
tural Experiment Station. Partial funding for the proj- 
ect was appropriated for the Minnesota Forest Bird 
Diversity Initiative by the Minnesota State Legisla- 
ture from the Environmental Trust Fund as recom- 
mended by the Legislative Commission on Minneso- 


ADKINS GIESE and CUTHBERT: WOODPECKER NEST TREE CHARACTERISTICS 


JTS 


ta Resources. We thank S. Gale, M. Friberg, and M. 
Knutson for assistance in the field study. G. Oehlert 
and C. Bingham provided statistical advice. We thank 
D. Andersen and G. Niemi for input and assistance 
with various aspects of the research. 


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THE CANADIAN FIELD-NATURALIST 


Vol 19 


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Received 27 February 2004 
Accepted 24 March 2005 


Detectability of Non-passerines Using “Pishing” in Eastern Ontario 
Woodlands 


J. RYAN ZIMMERLING 


University of Western Ontario, Department of Biology, London, Ontario N6A 5B7 Canada 
Current address: Bird Studies Canada, 115 Front Street, Port Rowan, Ontario NOE 1MO Canada 


Zimmerling, J. Ryan. 2005. Detectability of non-passerines using “pishing” in eastern Ontario woodlands. Canadian 
Field-Naturalist 119(3): 377-380. 


During spring and summer 1997, non-passerines were surveyed in three woodlots near Arnprior, Ontario, using standard 
point counts, and standard point counts combined with “pishing” (pishing involves the observer saying the words “pish pish 
pish pish” in a continuous series of short bursts). Of the 27 non-passerine species detected, 22 were recorded on more days 
using pishing as opposed to the standard point co.unt method. However, only three of these species were recorded on signif- 
icantly more days using pishing. Several woodpecker species approached more closely during point counts with pishing, 
which facilitated identification. In contrast, raptors and some other non-passerines that may have otherwise gone unnoticed 
were identified as they fled from the pishing sound. Hence, when the overall goal of research is to detect species richness or 
to gather presence and absence data in woodlands, point counts combined with pishing may increase detectability of some 


non-passerines. 


Key Words: breeding birds, non-passerines, pishing, point counts, woodlands, Arnprior, Ontario. 


Birds are surveyed by numerous methods includ- 
ing line transects, territory mapping, mark-recapture, 
point count, and playback recordings (Bibby et al. 
1992). The point count method relies upon the ability 
of the observer to correctly identify birds using both 
visual and auditory cues. For most observers, identi- 
fying birds by song is more difficult than is visual 
detection (Faanes and Bystrak 1981). In addition, some 
species of birds, especially non-passerines, are rarely 
heard during the breeding season and thus are usually 
only identified visually or by call-note; e.g., Sharp- 
shinned Hawk (Accipiter striatus). Moreover, songs 
or calls of some species are practically indistinguish- 
able from others. Many species of woodpeckers, for 
example, cannot be positively identified by drumming 
alone because of the high variation in resonance among 
drumming substrates (Robbins and Stallcup 1981). 
Numerous species of birds also are excellent mimics. 
The Blue Jay (Cyanocitta cristata) can imitate the call 
of the Broad-winged Hawk (Buteo platypterus). These 
difficulties, inherent in the aural detection of some 
bird species, have resulted in other surveying meth- 
ods being devised that augment detection probability. 

Playback recordings of bird songs and calls have 
been used primarily for augmenting more convention- 
al survey techniques. This method is primarily used 
for detecting nocturnal non-passerines and other birds 
that do not normally vocalize by song (Johnson and 
Brown 1981; Gunn et al. 2000). Playbacks of Red- 
shouldered Hawks (Buteo lineatus) are used during the 
breeding season to elicit a territorial response from 
this uncommon species (Badzinski 2003*). Although 
playbacks often are useful for attracting conspecifics 
closer to the observer, a better playback method would 


attract a number of different bird species. Many natu- 
ralists, bird photographers, and several researchers 
have used “pishing” (described by repeating the words 
“pish pish pish pish pish’”) to entice various species 
of birds to approach the observer (e.g., Emlen 1971; 
Wiedner et al. 1992; Runtz 1995; Porneluzi and 
Faaborg 1999; Sibley 2002; Prescott 2003*). In addi- 
tion, several long-term bird monitoring programs such 
as the Christmas Bird Count and the Ontario Breeding 
Bird Atlas also permit pishing to be used to gather 
species distribution data (Ontario Breeding Bird Atlas 
2001*). Interestingly, many European bird species 
appear unresponsive to pishing (Purdy 1998*; BBC 
2003*; but see Foppen et al. 2000). Some researchers 
believe that pishing simulates the sounds of a group 
of birds mobbing a predator while others believe that 
it is curiosity alone that attracts the birds to the sounds 
(Runtz 1995, Zimmer 2000). 

There are few studies that quantify the effective- 
ness of pishing. Lynch (1995) found that aural stimuli 
using playbacks, owl imitations, and pishing increased 
number of species detected. However, Lynch did not 
attempt to distinguish separate effects of the three 
stimuli tested. Zimmerling and Ankney (2000) found 
that, on average, 3.6 more passerine species were 
detected using pishing during spring point counts than 
point counts without pishing. Pishing has also been 
shown to be an effective technique for augmenting fall 
and winter point counts (MacDonald 2003*). Although 
primarily used for attracting passerines, it has been 
suggested that some hawks, owls, hummingbirds, and 
woodpeckers will also respond to pishing (Zimmer 
2000; BBC 2003*). To my knowledge, however, there 
are no data that examine the effectiveness of pishing 


eit | 


378 


with respect to detecting non-passerines during the 
breeding season. 

Therefore, the primary objective of this study was 
to determine if pishing in conjunction with the stan- 
dard point count method (hereafter referred to as 
pishing) would affect the number of non-passerine 
species detected relative to the standard point count 
method (hereafter referred to as standard method). 


Methods 
Study Area 

This study was conducted in three woodlots located 
near Arnprior, Ontario (45°26'N, 76°21'W). The wood- 
lots varied in size (approximately 0.75 km?, 2 km? 
and 2.25 km?), but each was a rectangular “island”, 
bordered on the north by the Ottawa River and non- 
maintained roads, fields, and residential areas on the 
other sides. A distance of 10 km separated the middle 
woodlot from woodlots to the west and east. Vegeta- 
tion was dominated by mature Sugar Maple (Acer 
saccharum) and American Beech (Fagus grandifolia) 
interspersed with White Pine (Pinus strobus). Herba- 
ceous plants dominated the understory vegetation. 
Each woodlot had at least one small marsh within its 
boundaries; and swamps were also present within the 
two largest woodlots. 


Bird Surveys 

Within each woodlot, two transect paths were 
flagged with plastic tape before surveying began. 
Each transect was divided into two lines that formed 
a 90° angle to each other. The first line of the first 
transect began in the southwest corner of the woodlot 
and bisected the northern boundary. The second line 
began 100 m to the east of the final point of the first 
line and terminated in the southeast corner of the 
woodlot. The second transect used the same line con- 
figuration as the first, but the orientation was reversed 
such that the first line of the second transect began in 
the northeast corner of the woodlot and bisected the 
southern boundary. The second line began 100 m to 
the west of the final point of the first line and termi- 
nated in the northwest corner of the woodlot. Num- 
ber of points per line varied with size of the woodlot: 
the smallest woodlot had 11 sampling points spaced 
>100 m apart, but lines for the two larger woodlots 
each had 15 sampling points spaced >100 m apart. 

Surveys were conducted from 28 April to 1 July 
1997 and began at sunrise (05:00 — 06:00 EST). Wind 
velocity, measured using a hand-held anemometer, 
and ambient air temperature, measured with a ther- 
mometer, were recorded before surveys began and 
after they were completed on each of the two lines. 
Surveys were terminated if wind velocity exceeded 
17 km/h or if precipitation occurred (Robbins 1981). 
Results of incomplete surveys were excluded from 
analysis. 

A different protocol was used to survey birds for 
each line on a given transect. Surveying both lines on 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


the same day using the different protocols controlled 
for variation in detectability that might have been 
attributable to change in weather from one day to the 
next. The first protocol employed standard point 
count methodology with birds surveyed for four min- 
utes at each point. Birds were counted if they were 
detected either aurally or visually within or below the 
forest canopy. Species and sex (whenever possible) 
were recorded for each individual detected and the 
mode of detection (visual or aural) that first allowed 
positive species identification was also recorded. Be- 
haviour of a detected bird towards the observer’s pres- 
ence was documented for each detection. 

During the second protocol, a recording of JRZ 
pishing was played. The recording only included the 
“pish pish pish pish pish” repertoire and did not in- 
clude squeaking, squeeling, owl imitations, or other 
noises. The pishing recording was analyzed using a 
spectrogram to ensure comparable quality and tempo- 
ral aspects with JRZ’s voice. The volume of the tape 
player was set before sampling at a volume similar to 
that of JRZ’s voice pishing. At each point, 30-second 
intervals of playing the pishing recording and listen- 
ing periods were implemented. The procedure was 
repeated three more times, with the speaker held 180° 
in the opposite direction each succeeding time such that 
birds on opposite sides of the line had equal detection 
probabilities. In total, two minutes were spent playing 
the recording and two minutes were spent in silence. 
Data on bird species were recorded as on the previous 
line with three minutes allotted for travel between sam- 
pling points. 

The order of protocols for each line of a transect 
was reversed every other day of surveying to control 
for variation in detectability that might be attributa- 
ble to time of day. Thus, each woodlot was surveyed 
for two consecutive days before the next woodlot 
was surveyed. After all woodlots had been surveyed 
using the first transect (six complete surveys), the 
second transect was surveyed (six complete surveys). 
Thus, 12 surveying days were required to survey all 
three woodlots using both transects. This was repeat- 
ed two more times for a total of 36 surveys. 


Data Analysis 

Chi-square analysis (PROC FREQ; SAS Institute 
2001) was used to determine if each bird species was 
detected more often using one of the two survey meth- 
ods. Only those bird species that were detected on five 
or more different days via one of the two methods were 
tested statistically. Birds that were detected between 
point count locations were excluded from the analysis. 


Results and Discussion 

Of the 15 non-passerine species wherein sample 
sizes were sufficiently large to test for statistical signif- 
icance, three species were detected on significantly 
more days using pishing as opposed to the standard 
point count method (Table 1). However, irrespective 


2005 


of statistical significance, 22 of the 27 non-passerine 
species detected were recorded on more days using 
pishing, whereas only three species were observed on 
more days using the standard method. Because sam- 
ple sizes were small, and encounters of some species 
were likely incidental (i.e., shorebirds), these results 
should be interpreted with caution. For example, shore- 
birds were unresponsive to pishing and most were 
detected on the same day and at the same location 
several hours after a severe thunderstorm presumably 
halted their migration activities. These results con- 
firm those by MacDonald (2003*) that suggested, in 
general, that passerine species seemed to approach 
more closely in response to pishing than did non- 
passerines, presumably because passerines mob more 
frequently. Indeed, both Lynch (1995) and Zimmerling 
and Ankney (2000) detected 19% more species dur- 
ing point counts using pishing than without, but those 
studies were limited to passerines during the breed- 
ing season. 

Interestingly, several woodpecker species were 
detected more often and approached more closely 
during point counts with pishing. For example, Hairy 
Woodpeckers (Picoides villosus) and Yellow-bellied 
Sapsuckers (Sphyrapicus varius) often approached 
within 15 m of the point count location when the pish- 
ing recording was played. Northern Flickers (Colaptes 


ZIMMERLING: USING “‘PISHING”’ IN EASTERN ONTARIO 


379 


auratus) were detected significantly more often using 
pishing than during the standard method but this 
species did not approach the point-count location. 
Other researchers (e.g., Zimmer 2000) have suggest- 
ed that woodpeckers are generally responsive to pish- 
ing, and therefore, it is not surprising that they have 
also been shown to mob predators (e.g., Gehlbach 
and Leverett 1995). 

Although pishing increased detectability by induc- 
ing some woodpeckers to approach point-count loca- 
tions more closely, detectability of other species was 
also increased due to birds fleeing (1.e., taking flight) 
from the pishing sound. For example, when pishing 
was used, Red-shouldered Hawks (Buteo lineatus) 
were recorded significantly more often as they flushed 
from perches. Other raptors, such as American Kestrels 
(Falco sparverius), Cooper’s Hawks (Accipiter co- 
operit), and Merlins (Falco columbarius) exhibited a 
similar response. Had these birds not fled from their 
perches, they might have gone unnoticed. This fleeing 
reaction by raptors is not surprising. Other studies 
have demonstrated that, as a result of harassment by 
mobbing passerines or advertisement of perception of 
the predator by the mobbers, predators usually leave 
(e.g., Bildstein 1982). Admittedly, for species that ex- 
hibited a flee response to pishing, any recorded, un- 
natural sound (e.g., clapping hands or shouting) proba- 


TABLE |. Number of days each non-passerine species was detected using the standard point-count method and pishing in 


Ontario woodlands, 1997. 


Common Name (Species name) 


Canada Goose (Branta canadensis) 

Wood Duck (Aix sponsa) 

Mallard (Anas platyrhynchos) 
Blue-winged Teal (Anas discors) 

Ruffed Grouse (Bonasa umbellus) 

Great Blue Heron (Ardea herodias) 
Osprey (Pandion haliaetus) 

Sharp-shinned Hawk (Accipiter striatus) 
Cooper’s Hawk (Accipiter cooperii) 
Red-shouldered Hawk (Buteo lineatus) 
American Kestrel (Falco sparverius) 
Merlin (Falco columbarius) 

Killdeer (Charadrius vociferus) 

Greater Yellowlegs (Tringa melanoleuca) 
Lesser Yellowlegs (Tringa flavipes) 
Spotted Sandpiper (Actitis macularia) 
Dunlin (Calidris alpina) 

Least Sandpiper (Calidris minutilla) 
Mourning Dove (Zenaida macroura) 
Barred Owl (Strix varia) 

Ruby-throated Hummingbird (Archilochus colubris) 
Belted Kingfisher (Ceryle alcyon) 
Yellow-bellied Sapsucker (Sphyrapicus varius) 
Downy Woodpecker (Picoides pubescens) 
Hairy Woodpecker (Picoides villosus) 
Northern Flicker (Colaptes auratus) 
Pileated Woodpecker (Dryocopus pileatus) 


* NS = no significant difference; nt = not tested (see text) 


Standard Pishing Pp? 


2 nt 
10 NS 


— 


OO, SON Cn Go) Sb) One SS SS So Orie SS oo "0 So 


— 
— 


NO — 
SONA AMAA OR RK RPK OANNK ABH KH ONY Co 
i=) 
or 


— 


bly would have induced the same behaviour. For 
example, on point counts using pishing, Great Blue 
Herons (Ardea herodias) were frequently detected as 
they flew from the emergent vegetation around swamps. 
Because better methods, such as playbacks, can be 
used to elicit a vocal response in some non-passer- 
ines, it is not recommended that pishing be used for 
the sole purpose of surveying these species. In addi- 
tion, Zimmerling and Ankney (2000) caution that 
pishing can confound relative abundance estimates of 
species that are particularly responsive to pishing. How- 
ever, when the goal of a study is to detect as many bird 
species as possible in a woodlot, regardless of taxon- 
omy, or to acquire presence/absence data for specific 
species, the results of this study suggest that pishing, 
when combined with standard point count methodol- 
ogy, may increase detectability of many bird species, 
including non-passerines. 


Acknowledgments 

I am grateful to Dave Ankney for his guidance and 
assistance throughout this project. I also thank Doug 
Tozer and Julie Zimmerling for reviewing earlier drafts 
of this manuscript. A. J. Erskine and two anonymous 
reviewers provided invaluable comments that greatly 
improved this manuscript. 


Documents Cited (marked * when cited) 

Badzinski, D. 2003. Red-shouldered Hawk and _ spring 
woodpecker surveys. Unpublished report by Bird Studies 
Canada for the Ontario Ministry of Natural Resources. 

BBC. [British Broadcasting Corporation] 2003. Calling all 
Birds. (20 March 2003) http://www.bbc.co.uk/nature/birds/ 
weeklyfeature/callingallbirds/index.html. 

MacDonald, M. 2003. Detectability of birds using point 
counts combined with pishing in woodlands during the 
fall and winter. Honour’s thesis. University of Western 
Ontario, London, Ontario. 

Ontario Breeding Bird Atlas. 2001. Guide for participants. 
Atlas Management Board, Federation of Ontario Natu- 
ralists, Don Mills. 

Prescott, D. R. 2003. The use of call playbacks for census- 
ing Loggerhead Shrikes in southern Alberta. Alberta 
Sustainable Resource Development, Fish and Wildlife 
Division, Alberta Species at Risk Report 67. 

Purdy, M. 1998. The art of pishing. Greater Akron Audubon 
Matters 24: 2. 


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census techniques. Academic Press, London. 

Bildstein, K. L. 1982. Responses of Northern Harriers to 
mobbing passerines. Journal of Field Ornithology 53: 7- 
14. 

Emlen, J. T. 1971. Population densities of birds derived 
from transect counts. Auk 88: 323-342. 


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Faanes, C. A.,and D. Bystrak. 1981. The role of observer 
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Foppen, R. B., J. P. Chardon, and W. Liefveld. 2000. 
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Gunn, J. S., A. Desrochers, M-A. Villard, J. Bourque, 
and J. [barzabal. 2000. Playbacks of mobbing calls of 
Black-capped Chickadees as a method to estimate repro- 
ductive activity of forest birds. Journal of Field Ornithol- 
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Johnson, R. R., and B. T. Brown. 1981. Playback recordings 
as a special avian censusing technique. Pages 68-75 in 
Estimating Numbers of Terrestrial Birds. Edited by C. J. 
Ralph and M. Scott. Studies in Avian Biology 6. 

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day, and aural stimuli on detectability of migratory and 
resident bird species in Quintana Roo, Mexico. Pages 1-6 
in Monitoring Bird Populations by Point Counts. Edited 
by C. J. Ralph, J. R. Sauer, and S. Droege. USDA Forest 
Service General Technical Report PSW-GTR-149. 

Porneluzi, P. A., and J. Faaborg. 1999. Season-long fecun- 
dity, survival, and viability of Ovenbirds in fragmented 
and unfragmented landscapes. Conservation Biology 
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Robbins, C. S. 1981. Bird activity levels related to weather. 
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Biology 6. 

Robbins, C. S., and R.W. Stallcup. 1981. Problems in sep- 
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Received 24 March 2004 
Accepted 7 March 2005 


Evidence of Range Expansion of Eastern Coyotes, Canis latrans, in 
Labrador 


Tony E. Cuusss! and FRANK R. PHILLIPS? 


'Department of National Defence, 5 Wing Goose Bay, Box 7002, Station A, Happy Valley-Goose Bay, Newfoundland and 
Labrador AOP 1SO Canada; E-mail: techubbs @cablelab.net 

*Department of Natural Resources, Government of Newfoundland and Labrador, Box 175, Station C, Happy Valley-Goose 
Bay, Newfoundland and Labrador, Newfoundland AOP 1CO Canada 


Chubbs, Tony E., and Frank R. Phillips. 2005. Evidence of range expansion of eastern Coyotes, Canis latrans, in Labrador. 
Canadian Field-Naturalist 119(3): 381-384. 


Eastern Coyotes were first documented in central Labrador in 1995 and have recently been recorded in coastal Labrador and 
at three additional locations in central and western Labrador. Here we document additional records indicating range expan- 
sion and the possibility of an established population. We also examine the future management of the species in Labrador 


and its possible effect on this northern ecosystem. 


Key Words: Coyote, Canis latrans, range, distribution, management, Labrador. 


During the middle of the nineteenth century, Coyotes 
(Canis latrans) expanded their distribution northward 
and eastward in North America (Nowak 1979; Pekins 
and Mautz 1988; Thurber and Peterson 1991; Lehman 
et al. 1991; Parker 1995). Coyotes were first recorded 
in western Québec in 1944, the Gaspé Peninsula of 
southeastern Québec in 1974 (Georges 1976; Créte 
and Desrosiers 1995), insular Newfoundland in 1985 
(Lariviere and Créte 1993; Parker 1995) and finally 
into Labrador in 1995 (Chubbs and Phillips 2002). 
Being adapted to a variety of food items and forest 
types and tolerant of human settlements has allowed 
the Coyote to expand its range. Here, we report on 
additional sightings, provide evidence for an estab- 
lished population and examine the consequences of 
future management of the species in Labrador and its 
possible effect on this northern ecosystem. 

The large canid and ungulate predator-prey system 
in Labrador is occupied by Wolves (Canis lupus), Cari- 
bou (Rangifer tarandus) and Moose (Alces alces). 
Black Bears (Ursus americanus) are also significant 
predators of these ungulates. This species complement 
separates northeastern Canada from most other regions 
in North America which support Coyote populations, 
including insular Newfoundland, where Wolves are 
absent. The expansion of Coyotes in eastern North 
America has been attributed to the clearing of forests 
for agriculture and timber that increased prey densities 
and the elimination of Wolves as major competitors 
(Lariviere and Créte 1993; Parker 1995). Hybridiza- 
tion of Coyotes with Wolves has resulted in eastern 
Coyotes being larger than western conspecifics (Créte 
and Desrosiers 1995) and recent information on ex- 
panding Coyote populations in eastern Québec also 
indicates that Coyotes appear to be larger in the eastern 
portion of their range (Thurber and Peterson 1991; 
Lariviére and Créte 1993; Peterson and Thurber 1993). 
Their larger size and the availability of White-tailed 


Deer (Odocoileus virginianus) were likely important 
in the successful colonization of Coyotes throughout 
northeastern North America (Larivére and Créte 1993). 
Coyotes prey on Woodland Caribou calves in Québec 
(Créte and Desrosiers 1995). In insular Newfound- 
land, Woodland Caribou appear to be the keystone 
ungulate prey species, which has enabled Coyotes to 
rapidly colonize much of the island (Wildlife Divi- 
sion, unpublished data). In Labrador, both woodland 
and migratory Caribou (George River Caribou Herd) 
may be important seasonal food sources (Chubbs and 
Phillips 2002). Although the northern forests are rec- 
ognized as sub-optimal habitat for Coyotes (Tremblay 
et al. 1998), they may persist at low densities, largely 
depending on the availability of Snowshoe Hare (Lepus 
americanus), mice and voles (Créte and Desrosiers 
1995; Samson and Créte 1997; Patterson et al. 2000; 
Crete et al’2001). 


Study Area and Methods 

Central Labrador lies within the Lake Melville-High 
Boreal Forest ecoregion and is dominated by Black 
Spruce (Picea mariana) with lesser amounts of Trem- 
bling Aspen (Populus tremuloides), White Birch (Betu- 
la papyrifera), Balsam Fir (Abies balsamea) and White 
Spruce (P. glauca) in river valleys (Meades 1990*). 
This region has the most favourable climate in Labra- 
dor with mean average daily temperatures ranging from 
-14 to -18°C in February and 13°C in July. Average 
annual precipitation is 1100 mm with an average snow- 
fall accumulation of 4 m (Meades 1990*). 

Coastal Labrador including Red Bay lies in the For- 
teau Barrens ecoregion and is primarily barrens vege- 
tated with Labrador Tea (Ledum groenlandicum) and 
Sheep Laurel (Kalmia angustifolia) with sparsely for- 
ested river valleys (Meades 1990*). This region has a 
high boreal climate with mean average daily temper- 
atures ranging from —9 to —12.5°C in February and 


a6! 


382 


12°C in July. Average annual precipitation is 1250 mm 
with an average snowfall accumulation of 3.5 to 4.5 m 
(Meades 1990*). 

We interviewed trappers in remote communities 
throughout Labrador and fur buyers to assess the oc- 
currence of Coyotes in the annual fur harvests. We also 
collected three carcasses, one harvested by trappers; 
the other killed on the highway. Local Conservation 
Officers reported one sighting near Goose Bay, Lab- 
rador. We found no other records of Coyotes harvested 
in other areas of Labrador. 


Results 

The first record of a Coyote in Labrador is that of 
an adult male, caught by a trapper on 14 January 
1995, along the Churchill River (53°17'N, 60°15'W) 
approximately 2 km south east of Happy Valley — 
Goose Bay (Chubbs and Phillips 2002). This speci- 
men was caught during a period of low trapping effort 
and which continues in Labrador today (Figure 1, 
Record I). 

Around 20 February 2003, an adult male Coyote was 
trapped near a landfill site (51°44.30'N, 56°25.00'W) 
just outside the town of Red Bay (Figure 1, Record 
IV). The skin (NFM MA-82.1) and skull (NFM MA- 
82.2) of this specimen were collected and positively 
identified as an adult male and are currently deposit- 
ed in the Newfoundland Museum of Natural History, 
St. John’s, Newfoundland and Labrador. 

On 19 January 2004 another adult male Coyote 
was picked up by Conservation Officers along the 
Trans Labrador Highway (53°02.22'N, 61°17.88'W) 
approximately 200 m west of Pena’s River and 75 km 
southwest of Goose Bay (Figure 1, Record II). The 
animal had apparently been killed by a vehicle. The 
carcass was largely intact although it had been par- 
tially scavenged. 

On 4 February 2004 on Lake Melville near Rabbit 
Island (53°23'N, 60°06'W), two Conservation Offi- 
cers on snowmobiles during daylight hours approached 
to within 10 m of a Coyote crossing the frozen lake 
(Figure 1, Record III). The officers, familiar with both 
Wolves and Coyotes, observed the animal for several 
minutes and confirmed that it was a Coyote. The Coy- 
ote was probably scavenging on fish offal left by ice- 
fishers in the area. 

In early December 2005, an adult male Coyote (M. 
Pritchett personal communication) was trapped near 
the landfill size 52°56.07'N, 66°51.24') approximately 
3 km east of the town of Wabush in western Labrador 
(Figure 1, Record V). The whole animal weight on a 
50 Ib scale was recorded at 45 Ib (20.4 kg) and the 
pelt was graded as XXL at the Northern American 
Fur Auction (J. Shouse, personal communication). 


Discussion 

The expansion of Coyotes in Labrador may have 
been influenced by the recent southern extension of 
the winter range of the George River Caribou Herd 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


during the last two decades. Caribou of the George 
River Herd have been recorded as far south as 52°10'N 
in central regions and 54°00'N near Cartwright along 
the coast of Labrador. Increased Moose densities in 
southern Labrador (Chubbs and Schaefer 1997) may 
provide additional food to support Coyotes through 
the winter. Coyotes are a highly adaptable species 
(Pilgrim et al. 1998) and are capable of surviving with 
a seasonally reduced prey base and harsh weather 
(Patterson et al. 1999). Coyotes are very efficient scav- 
engers and they may have expanded their range along 
a coastal route, scavenging on remains of abundant 
herds of Harp Seals (Pagophilus groenlandicus) and 
the less abundant Ringed (Pusa hispida) and Hooded 
seals (Crystophora cristata), and various fish and 
marine birds. Since 1992 the Trans Labrador Highway 
has provided an interior travel corridor between east- 
ern Québec, Wasbush, and Goose Bay. More recently 
a coastal highway from Québec to Red Bay and Cart- 
wright may provide an additional route for Coyotes 
to reach southeastern Labrador. 

The additional records reported here support the 
possibility of an established population in Labrador, 
as Coyotes are excellent colonists (Pilgrim et al. 1998) 
and coexist, thought at low densities, with Wolves in 
Alaska (Thurber and Peterson 1991), Alberta, British 
Columbia, Yukon and more recently Idaho (Pilgrim 
et al. 1998). No geographic barriers exist to discourage 
Coyotes from extending their range northward into 
Labrador. Delineation of Coyote range expansion in 
Labrador may have been delayed due to the absence 
of systematic surveys for carnivore species and because 
low fur prices in the last decade decreased trapping 
effort (only three trapped specimens over a ten-year 
period). It has been known for some time (Lehman et 
al. 1991; Pilgrim et al. 1998) that hybridization of Wolf 
and Coyote genotypes has occurred in Québec due to 
the rapid northeast progression of Coyotes. More re- 
cently it has also been shown that eastern Coyotes 
share a common evolutionary ancestry with the North 
American eastern Wolf (Wilson et al. 2000; Wilson et 
al. 2003). The larger size of eastern Coyotes may have 
contributed to their success in range expansion into 
Labrador. Eastern Coyotes make use of snow packed by 
human activities, especially snowmobile trails, facili- 
tating their travel and dispersal (Créte and Lariviére 
2003). Coyotes possibly maintain viable populations in 
northern forests, due to their high adaptability, great 
mobility, and low trapping effort (Trembley et al. 1998). 

Although a breeding record or incidence of an adult 
female Coyote has not yet been recorded in Labrador, 
the five records reported here, four of which were adult 
males, provide evidence of an established population. 
Labrador’s healthy Wolf population may be a control- 
ling mechanism, preventing a rapid population increase 
that has been observed in insular Newfoundland and 
elsewhere where Wolves are absent. Wolves are intoler- 
ant of Coyotes, killing them where both species overlap, 
permitting Coyotes to exist at low densities on the edge 


2005 CHUBBS AND PHILLIPS: RANGE EXPANSION OF COYOTES IN LABRADOR 383 


VA 


LABRADOR” 


/ 


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‘ se 2 ’ Se 
; ' 


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Coramunities 
Sightings 
Roads 
Proposed Road 
Rivers 


Trapping Locations 


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a 


Pena’s River s% 


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FicureE |. Location of Coyote (Canis latrans) records in Labrador. Sites I, IV, and V are trapping locations. Site II is the 
road-killed specimen and Site HI is an observation of a live Coyote. 


of established Wolf territories or within urban areas 
where Wolves rarely frequent (Parker 1995). 


Management Implications 

Since the time of European settlement of North 
America, Coyotes have been considered a predator that 
must be controlled. The Coyote reached the island of 
Newfoundland in 1985 and a trapping season was 
established in 1986. A trapping season for Coyotes was 
established in Labrador in 1995 shortly after the first 
confirmed record of the species in February of that year 
(Chubbs and Phillips 2002). Both regions allowed 
hunting in 2002. In areas where Coyotes and Wolves 
coexist they do so at reduced densities through spatial 
avoidance and changes in behaviour (Arjo and Pletscher 
1999; Switalski 2003). We speculate that although Coy- 
otes may not exist at high densities in Labrador due to 
the presence of Wolves, they may hinder the recovery 
of threatened Woodland Caribou populations in the area 
and we encourage studies to evaluate impacts. 


Acknowledgments 

Thanks are extended to trapper R. Moores for pro- 
viding the skull and skin of the Red Bay specimen 
and details of the capture location. Trapper J. Foote 


provided details of the speciman caught in Labrador 
west. We also extend thanks to Conservation Officers 
W. Lyall for details on the recovery of the road-killed 
Coyote, H. Yetman and B. Duffett for their observa- 
tions of the Coyote sighting near Rabbit Island, and 
M. Pritchett for details of the Labrador west specimen. 
M. Créte, N. P. P. Simon and G. Parker provided com- 
ments on earlier drafts of the manuscript. L. Elson 
drafted the figure. Funding for this publication was pro- 
vided through the Department of Natural Resources. 


Documents Cited (marked * in text) 

Meades, S. J. 1990. Natural regions of Newfoundland and 
Labrador. Protected Areas Association, St. John’s, New- 
foundland and Labrador. 103 pages. 


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Chubbs, T. E., and J. A. Schaefer. 1997. Population growth 
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Wilson, P. J., S. Grenwal, I. D. Lawford, J. N. M. Heal, A. 
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Received 7 September 2004 
Accepted 6 September 2005 


Consequences of Beaver, Castor canadensis, Flooding on a Small 
Shore Fen in Southwestern Quebec 


JOYCE M. REDDOCH and ALLAN H. REDDOCH 
548 Rivershore Crescent, Gloucester, Ontario KIJ 7Y7 Canada 


Reddoch, Joyce M., and Allan H. Reddoch. 2005. Consequences of Beaver, Castor canadensis, flooding on a small shore fen 
in southwestern Quebec. Canadian Field-Naturalist 119(3): 385-394. 


We describe the changes in a small peatland in Gatineau Park, Quebec, between 1878 and 2004. We correlate these changes with 
water levels and Beaver activity during this period. A shore fen persisted until 1980 under a regime of relatively stable water 
levels interspersed with episodes of somewhat higher water that killed encroaching trees. Significantly higher water levels in 
the early 1980s transformed the fen into a shallow water wetland containing a few partially-floating fragments of the fen. In 2003 
a beaver dam collapse lowered the water level dramatically, exposing the peat surface and making it available for colonization 
by existing aquatic and shoreline communities. If maintaining the fen had been part of the Park’s beaver control program, 
water levels should have been allowed to fluctuate in order to control tree growth but not permitted to rise high enough or for 
long enough to drown the fen. 


Key Words: Beaver, Castor canadensis, fen, flooding, floristics, shallow water wetland, shore fen, wetland succession, water 


levels, Quebec. 


The Precambrian uplands of eastern Ontario and 
western Quebec adjacent to the Ottawa River valley are 
dotted with many small lakes, a number of which sup- 
port fens along their shores. Although botanical explo- 
ration of the Ottawa area began in the 1860s, there 
appear to be few published floristic or ecological studies 
of these fens. A start on describing the calcareous fens 
within 50 km of the Parliament Buildings in Ottawa 
was made in the late 1970s (Reddoch 1979, 1984, 1989; 
Reddoch and Reddoch 1997). 

By the end of the 1920s Beaver populations in east- 
ern Canada and elsewhere had declined to low levels 
because of excessive exploitation. Thereafter, popula- 
tions began to rise again, partly the result of quotas, 
trap line restrictions and reintroductions of breeding 
animals (Banfield 1974; Peterson 1966) and partly 
because some human activities created new Beaver 
habitat (Ingle-Sidorowicz 1982). In Quebec, trapping 
was banned altogether between 1931 and 1941, and 
among the places where breeding pairs were introduced 
was Gatineau Park, across the Ottawa River from 
Ottawa, Ontario. By the end of the 1950s, Beavers were 
widespread and abundant in the park and adjacent areas, 
as was evident from the many new Beaver dams and 
subsequent flooding of low-lying areas. By the late 
1970s, a program to control water levels in Beaver 
ponds was initiated in Gatineau Park in order to ensure 
public safety and protect man-made structures (Nation- 
al Capital Commission 2004*). Since then, there have 
been ten aerial surveys to locate and census the popula- 
tion. The most recent survey, in 2002 — 2003, found 283 
active colonies, an average of 0.78 colonies per square 
kilometre (I. Beaudoin-Roy, personal communication). 
In Gatineau Park, Beavers are present in an area for a 
year or more and then are absent for another period of 


time. They seem to move about more during times of 
high precipitation (M. Leclair, private contractor, per- 
sonal communication). In northeastern North America, 
Beavers frequently inhabit peatlands, especially where 
there is open water. Their flood waters can modify or 
kill the vegetation on grounded parts of peatlands while 
having varying effects on those parts of the peatland 
that are floating (Damman and French 1987; Mitchell 
and Niering 1993; Muir-Hotaling c. 2001*; Rebertus 
1986; Schwintzer and Williams 1974; personal obser- 
vation). 

We have been doing botanical work in Gatineau Park 
since the late 1960s, concentrating especially on long- 
term studies of some native orchid colonies (Reddoch 
and Reddoch 1997). In the spring of 2003 we noticed 
that the water level of Black Lake suddenly had 
dropped a metre, exposing an expanse of peat in the 
southeastern corner of the lake where we had observed 
a fen in the 1960s and 1970s. We report here the 
changes in the wetland that we have recorded on a spo- 
radic basis between 1967 and 2004. From literature ref- 
erences, herbarium collections, air photographs and 
wood samples, we deduce the various stages of this 
wetland since 1878, a total of 127 years. We correlate 
these observations with the water levels and Beaver 
activity during this period and consider what manage- 
ment strategies would have been necessary to preserve 
the fen habitat. 


Study Site 

The study site is about 15 km west-northwest of 
the Parliament Buildings in Ottawa, Ontario, at the 
edge of Black Lake (45°29'29"N, 75°51'46"W) in 
Gatineau Park. The park, which is in the Quebec por- 
tion of the National Capital Region, is 363 km? in 


385 


386 


area; it is owned and managed by the National Capi- 
tal Commission. 

The climate of the region is continental, with warm, 
moist summers and cold, snowy winters. The mean 
daily maximum and minimum July temperatures at 
Ottawa are 26.5°C and 15.4°C, while the correspon- 
ding January temperatures are -6.1°C and -15.3°C. 
Mean annual precipitation is 943 mm, of which one 
quarter falls as snow. Mean precipitation for the month 
of July is 90.6 mm; the mean frost-free period is 159 
days, extending from the beginning of May to early 
October (Environment Canada 2004*). At Black Lake, 
300 m above mean sea level and about 200 m higher 
than Ottawa, the temperatures are slightly cooler, the 
precipitation is slightly higher and the mean frost- 
free period is slightly shorter (Crowe 1984; Environ- 
ment Canada 2004*). 

Black Lake is on King Mountain, part of the Eardley 
Escarpment that defines the southwestern exposure of 
the Canadian Shield in this area. The lake is situated in 
an east-west trough along a fault line between two 
ridges rising 50-60 m above it. The bedrock is Precam- 
brian diopsidic gneiss interspersed with granite peg- 
matite (Hogarth 1962, 1970); it is exposed or shallowly 
covered with glacial till. The lake is surrounded by 
forests more than 80 years old that are characteristic of 
the Middle Ottawa Section of the Great Lakes — St. 
Lawrence Forest Region of which they are a part 
(Lopoukhine 1974; Rowe 1972). Much of what is now 
Gatineau Park was logged and subsequently burned 
over in the 1800s and early 1900s (Mott and Farley-Gill 
1981). The north-facing slope to the south, and adjacent 
eastern slope, are covered predominantly with Eastern 
Hemlock (Tsuga canadensis (L). Carriere) accompa- 
nied by a few scattered individuals of Eastern White 
Pine (Pinus strobus L.), Eastern White Cedar (Thuja 
occidentalis L.), Yellow Birch (Betula alleghaniensis 
Britton),White Birch (B. papyrifera Marshall) and Red 
Maple (Acer rubrum L.), while the forest on the south- 
facing slope farther away to the north is mainly decidu- 
ous with Sugar Maple (Acer saccharum Marshall) and 
Red Oak (Quercus rubra L.) co-dominant. 

Black Lake is about 2 ha in area; it is 150 m across 
and has a maximum depth of about 8 m. Much of the 
shoreline is bedrock. Measurements made at the 2-m 
depth in the lake on 19 July 1982 (Eco-recherches 
Inc. 1983*) were as follows: water temperature 21.5°C, 
pH 7.1, dissolved oxygen 7.9 ppm, carbon dioxide 
6 mg/L, alkalinity 10 mg/L, total phosphates 0.04 mg/L 
and sulphates 14 mg/L. The only sources of water are 
rainfall and runoff from the surrounding slopes, which 
constitutes a catchment area of about 8 ha. The out- 
let, and consequently the location of the Beaver dam 
that affects water levels, is in the southwestern corner 
of the lake. The subject wetland, 0.12 ha in area, is 
situated in the southeastern corner, adjacent to the 
north-facing, 50°slope (Figure 1). This slope pro- 
vides a cool microclimate for the wetland by shading 
it for much of the year, except in the summer. The ice 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


forms first in this part of the lake in the fall and thaws 
last here in the spring. 


Materials and Methods 

Air photos were examined at the National Air Photo 
Library in Ottawa. Useful images were available for 
1925, 1928, 1935, 1938, 1944, 1948, 1951, 1966, 1971, 
1988, 1994, 1999, 2001 and 2002. Some photos were 
not useful because they were taken in the spring, winter 
or autumn when the sun was low and the north-facing 
slope cast a heavy shadow that obliterated details of the 
wetland and the outlet stream. Scales ranged from 
1:6000 to 1:16 000. 

Our field notes, as well as our photographs taken at 
various times between 1967 and the present, provided 
considerable information. Field work in 2003 and 2004 
consisted of identifying the flora on the newly exposed 
expanse of peat and mapping the area with a 100’ steel 
tape and compass. Inaccessible parts of the site were 
examined with binoculars. Water levels were estimated 
from the shapes of the lake in the air photos and, in the 
field, from the positions of the water in relation to the 
high-water mark on the steep bedrock face adjacent to 
the wetland. 

Information about the historical tree inventory of the 
fen was obtained from old stumps and fallen logs by 
wood identification techniques (Hoadley 1990; Panshin 
and de Zeeuw 1980). Twenty-six wood samples, a few 
centimetres in size, were collected and determined to 
genus level or better. The effects of water immersion 
and decay caused some difficulties in the identifica- 
tions. Another difficulty arose with Thuja occidental- 
is samples because these trees were the stunted form 
found in fens that are not considered in most commer- 
cial timber identification work. This form is represented 
by a dead T: occidentalis tree we had collected in anoth- 
er Ottawa area fen (Reddoch 1984). When it died, it was 
43 years old, Im tall and 3 cm in maximum diameter. 
It was identified partly by its remnants of fibrous inner 
bark. Like some samples from Black Lake, its wood 
was hard with annual rings as narrow as 0.1 mm. In 
contrast, a sample of T: occidentalis from a stump on 
the shoreline of Black Lake near the wetland was quite 
normal with annual rings about 4 mm wide. It was also 
difficult to distinguish Betula from Alnus because the 
set of diagnostic characters sometimes gave conflicting 
results, likely the consequence of a nutrient-poor habi- 
tat. However, in the case of stumps and trunks more 
than 20 cm in diameter, A/nus can be excluded. In 
some cases Betula papyrifera was readily identified 
by remnants of bark. 

Vascular plants collected during this study have 
been deposited in the herbaria of the Canadian Muse- 
um of Nature (CAN) and Agriculture Canada (DAO) 
in Ottawa. Grasses were identified by S. J. Darbyshire 
at DAO in accordance with Dore and McNeill (1980). 
Mosses collected in 1978 were identified by R. R. 
Ireland and deposited in Canadian Museum of Nature 
Moss collection (CANM). Vascular plant (except grass- 


2005 REDDOCH and REDDOCH: BEAVER FLOODING ON A SMALL SHORE FEN 387 


* 


% 8, sa Pe y bes 
CO TE Saal wee MLSE A 


FIGURE I. The shore fen at Black Lake as viewed from the slope to the north on 27 July 1971. Photograph by Joyce M. Reddoch. 


388 


es) and bryophyte nomenclature follow Newmaster et 
al. (1998). The principal depositories for local collec- 
tions, CAN and DAO, were searched for possible 
previous Black Lake collections by looking for species 
that are characteristic of peatlands in this area. No 
additional species were found. The McGill University 
Herbarium (MTMG) was examined for Ottawa District 
orchid collections. 

Where possible, the terminology used for wetlands is 
from The Canadian Wetland Classification System 
(National Wetlands Working Group 1997) and Crum 
(1988). In particular, shallow water wetlands have 
“free surface water up to 2 m deep, present for all or 
most of the year, with less than 25% of the surface 
water area occluded by standing emergent or woody 
plants. Submerged or floating aquatic plants usually 
dominate the vegetation” (National Wetlands Working 
Group 1997). Fens are “grass-, sedge-, or reed-domi- 
nated peatland[s], often with some shrub growth or a 
scant tree cover, developed under the influence of min- 
eral-rich, aerated water at or near the surface. ...” (Crum 
1988). 


Results 

1878-1889. On 8 July 1878, James Fletcher collect- 
ed Pogonia ophioglossoides (DAO 767100) and Platan- 
thera clavellata (Michx.) Luer (DAO 267278, MTMG 
47571; sub Habenaria tridentata Hook.) in what he 
described in his Flora Ottawaensis (Fletcher 1893) as “a 
tiny bog at Black Lake, high on King’s [sic] Mountain, 
Chelsea, P.Q.” In earlier parts of the same publication 
(Fletcher 1889a, 1889b) he listed Solidago uliginosa 
Nutt. from a “swamp on King’s [sic] Mountain” and 
Utricularia cornuta Michx. from “a small swamp at 
Black Lake Kingsmere”’. (In his writings and on herbar- 
ium labels Fletcher used “swamp” and “bog” inter- 
changeably. Chelsea [now called Old Chelsea] and 
Kingsmere were nearby hamlets at this time.) It is likely 
that all four species were recorded from the same locali- 
ty; these species taken together suggest an open fen 
habitat. 


1925-1938. The air photos taken in 1925 and 1928 
show Black Lake surrounded by forest that extended to 
the water’s edge. By contrast, water levels in 1935 and 
1938 had risen somewhat and the trees along the north- 
ern and northeastern shores were no longer present. A 
number of fallen trunks were visible in the water where 
the trees had previously been. (In 2003 the stumps of 
these trees were once again exposed. All of the trees we 
sampled were Betula papyrifera.). In the southeastern 
corner of the lake, the triangular area of the wetland was 
covered with moderately spaced, moderate-size trees, 
mostly deciduous, except for a wide band along the lake 
edge that appears to be open fen. The width of the band 
was calculated from the 1928 air photo to be about 3 m. 


1938-1951. From 1938 to 1944 the lake level ap- 
pears to have dropped somewhat. However, in 1948 
the shape of the lake indicates a somewhat higher water 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


level and there appears to be a band of water between 
the wetland surface and the shore. By 1951 the band 
had disappeared but the water was still relatively high. 
Trees were no longer evident in the wetland, but the fen 
mat remained. There was now a distinct treeless shore- 
line around the rest of the lake about 10 m wide show- 
ing that sometime during this period water levels had 
been even higher. 


1966-1978. In the late 1950s a scenic parkway was 
built across the northern slope within 50 m of Black 
Lake. Its embankment encroaches slightly on a small 
section of the high-water north shore. Its design seems 
to have allowed for natural drainage into the lake, and 
any impact on the lake, other than visual, is not evi- 
dent. The parkway is closed in the winter. The air pho- 
tos of 1966 and 1971 indicate that the lake had returned 
to the low levels of the 1920s. A 1966 air photo is the 
first one to show evidence of Beavers: a lodge in the 
northwest corner of the lake. Aiken and Gillett (1974) 
reported the presence of a Beaver dam in 1971. We 
visited and photographed the fen from time to time dur- 
ing this period and in 1978 made a species list and col- 
lected a few specimens. [See Reddoch and Reddoch 
(1987) for a photograph of the fen in 1978.] The wet- 
land consisted of an open, poor fen mat edged along the 
lake with a semi-floating shrub margin. The poor fen 
mat (Sphagnum lawn) was dominated by Sphagnum 
magellanicum and supported Thelypteris palustris, 
Pogonia ophioglossoides, Drosera rotundifolia and Tri- 
adenum virginicum. Some plants of Carex cryptolepis 
Mack., Carex sp., Juncus brevicaudatus, Iris versicolor, 
Cicuta bulbifera, Lycopus uniflorus, Aster puniceus 
L., Bidens sp., Euthamia graminifolia and Solidago sp. 
were also present. There were scattered shrubs of 
Chamaedaphne calyculata, as well as some saplings of 
Alnus incana ssp. rugosa and young trees of Betula 
papyrifera that had grown to a height of 4 m by 1978. 
A few plants of Calla palustris L. grew in water-filled 
depressions in the mat. A mid-summer paper pH read- 
ing of the surface fen water in 1977 was 5.5 The semi- 
floating mat at the lake/fen interface (shrub fringe) 
consisted mainly of Chamaedaphne calyculata, which 
was accompanied by the mosses Sphagnum magellan- 
icum, S. teres, Calliergon stramineum (Brid.) Kindb. 
and Campylium polygamum (Schimp. in B.S.G.) C. 
Jens. The shape of the fen/lake interface was essentially 
unchanged from that of 1925. There was a narrow 
aquatic vegetation zone around most of the lake shore 
but not in front of the fen, presumably because the 
water was too deep there. Nuphar variegata was visual- 
ly the dominant species. (In 1971 Aiken and Gillett 
(1974) identified 19 species of aquatic vascular plants 
around the lake.) In the late 1960s and early 1970s, 
there were a few plants of the orchid Platanthera 
clavellata, which had been collected by Fletcher in 
1878, on the lake shore to the north of the fen, but we 
did not find any plants in the fen. In 1978 we noticed 
that the fen had become somewhat wetter than previ- 


2005 


ously and a stash of deciduous branches along the lake 
edge of the fen was evidence of Beaver activity. 


1983-2002. In 1983 the Sphagnum lawn had been 
replaced by open water dotted with the still-standing 
dead Betula papyrifera trees. Much of the Chamae- 
daphne calyculata shrub fringe and adjacent Sphag- 
num lawn remained intact as a semi-floating band 
2-3 m wide, although many of the shrubs were dying. 
A large Beaver lodge was prominent in the shrub 
fringe. The water level in 1983 was at least 30 cm 
above that of 1978, and by 1988 was a metre higher 
than in the 1970s. An aquatic community similar to 
that around the rest of the lake and consisting most 
prominently of Nuphar variegata, Sparganium amer- 
icanum and Carex sp. became established in the area 
of the former grounded Sphagnum lawn. By 1990 only 
stubs remained of the B. papyrifera trees; the trunks 
likely had provided building materials for the two 
additional Beaver lodges that had appeared by that 
time. The emergent parts of some larger fallen trunks 
provided sunning places for Painted Turtles (Chryse- 
mys picta) and Northern Leopard Frogs (Rana pipi- 
ens). In 1992 we photographed a Blanding’s Turtle 
(Emydoidea blandingi) there, a species that is extreme- 
ly rare (S1) in Quebec (Société de la faune et des parcs 
2004*). 

With continuing higher water, the former Chamae- 
daphne fringe became a line of low-lying islands, which 
can be seen in air photos as an irregular necklace along 
the line of the former fen margin and at the edge of the 
underlying peat. Over the years, the islands slowly have 
continued to fragment and disintegrate. The fragments 
have moved shoreward away from their original loca- 
tions, likely the result of buffeting by wave and ice 
action driven by the prevailing westerly winds. The 
dominant vegetation on the islands changed from 
Chamaedaphne calyculata to Carex lasiocarpa, al- 
though the former was still well-represented. In 2002, 
July and August were hotter than normal and among 
the driest on record (Environment Canada 2004*). As 
a result, the water level of the lake dropped about 0.75 m 
and three islands that were close to the shores, one at the 
north end and two at the south end, were left beached. 

The northern island measured 0.9 m x 1.30 m; the 
depth of the peat base as it sat compressed by its own 
weight was about 30 cm. All of the species included in 
the 1978 description of the Sphagnum lawn (above) 
were present, except for Cicuta bulbifera, Aster puni- 
ceus, the Solidago species and Betula papyrifera. There 
was, in addition, one plant each of five other species: 
Typha latifolia, Carex pseudo-cyperus, Viola mac- 
loskeyi ssp. pallens, Lysimachia thyrsifolia and Eupa- 
torium perfoliatum. Alnus incana was represented by 
one seedling 15 cm high. The dominant moss, Sphag- 
num magellanicum, was accompanied by S. teres and 
Campylium sp. [See Appendix for complete species list. 
A close-up photograph of this island with some of its 
eight flowering stems of Pogonia ophioglossoides is 


REDDOCH and REDDOCH: BEAVER FLOODING ON A SMALL SHORE FEN 


389 


shown in St. George (2002).] By contrast, the two peat 
islands that were stranded on the south shore had a lim- 
ited number of species, the dominant cover being Carex 
sp., accompanied by a few plants of Thelypteris palustris 
and Chamaedaphne calyculata, as well as some Sphag- 
num teres. The dimensions of these two islands were 
1.7 m X 2.2 m and 0.85 m x 0.40 m. These three is- 
lands at the northern and southern limits of the island 
chain represent the extremes in the gradient of the plant 
compositions of the islands. The farther north the is- 
lands were situated, the more species they supported. 


2003-2004. In late May of 2003, the lake was at its 
highest level, as was usual after snow melt. Two weeks 
later the water level had dropped about a metre because 
the Beaver dam had given way. Great volumes of water 
had cascaded down the escarpment onto the lowlands 
200 m below (M. Leclair, personal communication). 
Because there were no Beavers in the lake in 2002, the 
dam had not been kept in good repair. A device had 
been installed in the dam that year to maintain a rela- 
tively constant water level (M. Leclair, personal com- 
munication) but apparently was not functioning ade- 
quately in the spring of 2003. 

As a result of the drop in water level, the remaining 
islands settled down onto the peat substrate of the for- 
mer fen. One fragment about a metre square was strand- 
ed on top of a large, previously submerged stump and a 
smaller piece came to rest on a fallen tree trunk. The 
substrate was relatively homogeneous, unconsolidated 
peat, with a shallow surface gradient upward from the 
lake. Throughout the summer the peat remained saturat- 
ed. The central area was covered with shallow pools, 
while the inland borders were moist. Overall there was 
a related gradient of gradually decreasing moisture from 
the lake edge to the inland shore of the former wetland. 
The peat surface was dotted with stubs and stumps 
rooted in the peat and corresponding-sized trunks lying 
on and in the peat. There were many Betula papyrifera 
stubs of 6-10 cm diameter, as well as a few Alnus 
incana stubs of similar size. Their presence is consistent 
with our 1978 observations. Also revealed were a num- 
ber of B. papyrifera stumps in the range of 30-45 cm 
basal diameter and fallen trunks of related diameters, 
some still retaining their bark. In addition there were a 
few similarly sized stumps of Pinus strobus and Thuja 
occidentalis, and trunks of the former. At one place, 
8 m from the shore, we inserted a 1.8 m probe its full 
length into the peat, encountering increasing resistance 
but not touching bottom. At another place, two metres 
from the shore, the probe descended one metre into the 
peat. In other places, the probe encountered large rocks 
like those that occur plentifully all along the base of the 
slope, as well as some softer objects that possibly were 
previous generations of trunks and stumps, all less than 
a metre below the surface. 

Along the northeastern and northern shores of the 
lake beyond the wetland, many more large Betula pap- 
yrifera stumps were exposed by the drop in water level. 


390 


They likely were the remains of the trees shown on the 
air photos lining the lake shore in the 1920s. Their posi- 
tions indicate that the drop in 2003 brought the water 
level back almost to the low level of the 1920s. 

As the summer progressed, vegetation developed 
quickly on the exposed peat. Two different but overlap- 
ping communities could be distinguished. A Sparga- 
nium—Sagittaria community was established closer to 
the lake, around and in shallow ponds where the water 
table was close to or at the surface (shallow pool zone). 
A community of moist to wet habitat, mostly weedy 
species formed on the broad band of saturated to moist 
peat between the previous zone and the upland shore 
(upper shore zone). (The Appendix lists the species that 
were recorded from each zone.) During the summer, the 
islands stranded in previous years on the north and 
south shores continued to dry out and disintegrate. 

In 2004 the water level continued at the 2003 level. 
The plant communities of the previous year became 
increasingly established. Only one new species was 
found, a few plants of Aster lanceolatus Willd. in the 
upper shore zone. Several additional species that were 
present in the upper shore zone in 2003 had invaded the 
northern beached peat island: Euphrasia stricta, Bidens 
connata, Euthamia graminifolia, Solidago canadensis 
and Dulichium arundinaceum. The base of this island 
was in contact with the lake water in the late summer 
and fall. All of the peat islands have disintegrated to 
the extent that they seem unlikely to survive for more 
than a few years. The Beavers built yet another lodge, 
this one over a peat island near the north end of the 
former fen. Thus, since 1978, Beavers have built five 
lodges around the lake, four on the former fen and one 
in the northwestern corner of the lake (at a different 
location from that shown on the 1966 and 1971 air 
photos). In 2004 two of the lodges showed evidence 
of occupation, the one that was built this year and the 
one across the lake. The seasonal variation in water 
level during these two years of relatively normal rainfall 
and temperatures was in the range of 20 cm, except for 
the record 14 cm rainfall from the remnant of Hurri- 
cane Frances, which briefly raised the water level by 
about 24 cm in September 2004. 


Discussion 

The level of the water in Black Lake appears to have been 
the most important factor in determining the nature of 
the wetland over the past century and a quarter, and pre- 
sumably for much longer. In the 1920s the water level 
was relatively low. Judging by the presence of large 
trees along the shores and in the wetland, we deduce 
that it had been low for many decades. There was an 
episode of fluctuating higher water levels in the 1930s, 
1940s, and early 1950s followed by a return to the for- 
mer low level. This level was maintained until the end 
of the 1970s. From the early 1980s to 2003 there was 
a period of even higher water than before. This second 
rise in lake level was the work of Beavers, and it may 
be that Beavers contributed to the high water of the 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


1930s and 1940s, perhaps in conjunction with other 
natural causes. The later increase in water level was of 
the order of one metre and we estimate that the earlier 
increase at its peak was about half that. In 2003 the 
water level dropped almost to the low level of the 1920s. 

Until the late 1940s, water levels remained low 
enough to support a peatland consisting of a lacustrine 
swamp with a band of open fen along the lake edge. 
The trees of the swamp appear in air photos to have 
been a moderate size in 1948. This size correlates with 
the 30-45 cm basal diameter stumps and corresponding 
trunks of Betula papyrifera that were exposed when the 
water receded in 2003. The few Pinus strobus and 
Thuja occidentalis stumps and trunks may be from the 
same generation of trees as the B. papyrifera. The trees 
that grew in the central part of the wetland show evi- 
dence of stunted growth, reflecting the poor growing 
conditions of the peatland. Betula papyrifera is a com- 
mon colonizer of open sites on a variety of soils, includ- 
ing wetlands (personal observation), and is often found 
in pure stands (Farrar 1995). Thus it is likely that the 
presence of B. papyrifera trees of an even age is evi- 
dence of a previous disturbance in the late nineteenth 
century. 

By 1951 the water evidently had risen high enough 
and lasted long enough to kill the trees in the swamp, 
but appears not to have seriously affected the open fen. 
Over the next decade the water receded to the former 
low level, perhaps because a newly built Beaver dam 
was left unattended and consequently deteriorated after 
the Beavers were trapped out or had moved on. By the 
1960s, a new generation of Betula papyrifera and Alnus 
incana began to grow in the area of the former 
swamp, which by then had developed into a Sphagnum 
lawn. The open fen was a semi-floating Chamaedaphne 
shrub fringe between the Sphagnum lawn and the lake. 
The peatland supported more than 18 species of vas- 
cular plants in 12 families. Most studies of the impact 
of higher water levels on peatlands have been made on 
bogs and have some parallels to the present work. 
For example, Schwintzer (1979) has described how 
moderately raised water levels killed the trees in a treed 
bog in Michigan. The resultant increased light on the 
bog mat enhanced the development of Sphagnum 
species and Chamaedaphne. 

Beavers were present in the lake in the 1960s and the 
impact of their increasingly high dam at the outlet of 
the lake began to affect the fen in the late 1970s. 
Because the Sphagnum lawn was grounded, rising 
water killed all of the vegetation, including the trees, 
within a few years. The area of the drowned fen was 
replaced by a Nuphar—Sparganium shallow water wet- 
land that lasted until the spring of 2003. The shallow 
water wetland essentially was an extension over the 
peat substrate of the aquatic zone already present around 
the other margins of the lake (Aiken and Gillett 1974). 

The original shrub fringe stayed afloat as the water 
rose but fragmented into a line of islands, which have 
disintegrated over time. Some of the Chamaedaphne 


2005 


shrubs died and Carex lasiocarpa became the co-domi- 
nant cover, perhaps because there was a change in the 
water regime. The composition of the vegetation com- 
munity on the islands also changed to form a gradient 
from south to north, the southern islands supporting a 
restricted number of species, while the composition 
of the northern islands resembled that of the previous 
Sphagnum lawn (see Appendix). The occurrence of this 
gradient perhaps can be correlated with the amount of 
sunlight available through the growing season, the 
southern islands being increasingly more in the shadow 
of the slope above them. For instance, from 1980 until 
the present, the heliophilous orchid Pogonia ophioglos- 
soides has been present predominantly on the northern 
islands. Assuming continuity of fen habitat from 1878 
to the present, this is the longest recorded colony of 
orchids in the Ottawa District (Reddoch and Reddoch 
1997): 

When the water level dropped a metre in the spring 
of 2003 as a result of the Beaver dam giving way, the 
peat generated over the years or centuries by fen vegeta- 
tion lay exposed. For the first time, emergent and other 
plants in the seed bank were given the opportunity to 
germinate. The vegetation that became established on 
the peat in the first summer consisted of essentially two 
zones: a Sparganium-Sagittaria community in the 
saturated peat and shallow pools closer to the lake, 
and an upper shore zone of moist to wet habitat species 
around the inland edge. A total of 39 species in 21 fam- 
ilies made up the vegetation communities in the two 
zones. Four of the 30 species (13%) in the upper shore 
zone were aliens (see Appendix). That there is spa- 
cial separation of species in relation to water level is 
well known (see, e.g., Odland and del Moral 2002). 
The species in the two zones are characteristic of these 
habitats in the Ottawa area. Most of the species in the 
upper shore zone were also present on the adjacent east- 
er shore of Black Lake. They included only two of the 
five most abundant species listed by Le Page and 
Keddy (1998) as buried seeds in nearby Beaver ponds 
in Gatineau Park. Of the total of 37 species that LePage 
and Keddy (1998) discovered in Beaver pond sediments 
in the park, only 11 species in all were present in the 
upper shore zone at Black Lake. The shallow pool zone 
included about eight of the 16 aquatic species that were 
identified in 1971 by Aitken and Gillett (1974) from the 
margins of the lake. 

As Crum (1988) has pointed out, it is difficult to esti- 
mate the rate of peat accumulation. He suggested that 
peatlands in the area of the upper Great Lakes can take 
100 to 900 years to produce 30 cm of peat. Using these 
numbers, one can deduce that the depth of the peat in 
the Black Lake wetland, which is at least 1.8 m deep, 
may have taken between 600 and several thousand 
years to develop. An accumulation rate consistent with 
Crum was obtained from a core taken in the Mer 
Bleue peatland in the Ottawa Valley 30 km to the south- 
east (Camfield 1969). Unlike the Mer Bleue, the 
Gatineau Park area where Black Lake is located was not 


REDDOCH and REDDOCH: BEAVER FLOODING ON A SMALL SHORE FEN 


flooded by the Champlain Sea but has been open since 
the Wisconsin glaciation retreated about 12 000 before 
present (Romanelli 1975). 

Mott and Farley-Gill (1981) state that the almost 
unchanged forest composition in the Gatineau Park area 
over the past 3200 years indicates that there have been 
only minor variations in the climate during that time. 
The only major changes in the forest composition have 
occurred since logging began in the area about 175 
years ago. It is not known what impact, if any, logging 
has had on the Black Lake wetland nor whether the 
trees in the wetland have ever been logged. A few of the 
larger stumps have flat tops, but the condition of the 
wood in all of them precludes any definite conclusions. 
It is possible that any dead trunks still standing in the 
mid-1960s were cut down at the same time that the 
adjacent eastern slope was cleared of undergrowth, 
presumably to improve the view from the newly built 
parking lot beyond the eastern slope. 

It is recognized that Beaver flooding in certain cir- 
cumstances can increase species richness and landscape 
diversity. In a forested region such as the Adirondacks, 
New York, ecosystem engineering by Beavers has 
introduced wetlands into the landscape (Wright et al. 
2002). On the other hand, at Black Lake, Beaver flood- 
ing destroyed the shore fen that had made a unique 
contribution to the biological diversity of Gatineau 
Park (Gillett and Catling 1983). The fen has been re- 
placed, over the short term at least, by plant species 
already well represented in the lake’s aquatic and 
shoreline flora. Fens are the product of centuries of 
development and cannot quickly redevelop. 

The provincially extremely rare Blanding’s Turtle 
was present in the shallow water wetland phase at 
Black Lake in 1994. Since this elusive turtle is known 
to occur in (and move between) various types of wet- 
lands, including peatlands (Bider and Matte 1996; 
Cook 1981, 1984; McMurray 1984*; Desroches and 
Rodrigue 2004), it is not known how the changes in 
Black Lake over the past several decades have affected 
its suitability for this species. This turtle is known in 
the province only from a few reports in southwestern 
Quebec (McMurray 1984*; Bider and Matte 1996; 
Desroches and Rodrigue 2004). 

Preserving the fen at Black Lake would have re- 
quired maintaining historic low water levels. Every 
few decades a short period of higher water (about half 
a metre higher) would have been required to kill the 
trees. If the preservation of existing natural communi- 
ties in Gatineau Park had been a criterion when the 
objectives for managing Beavers were drawn up, much 
more careful attention would have had to be given to the 
impact of changing water levels on these communities. 


Acknowledgments 

We are grateful to Francis R. Cook, Researcher 
Emeritus, Canadian Museum of Nature, for information 
and valuable comments on the Blanding’s Turtle in 
Quebec, as well as confirming our identification of the 


302 


Blanding’s Turtle that we photographed at Black Lake; 
to Stephen J. Darbyshire, Biologist, Agriculture and 
Agri-food Canada, for identifying the grasses; to Robert 
R. Ireland, former Curator, Canadian Museum of 
Nature, for identifying the 1978 mosses; and to Donna 
Naughton, Research Assistant, Research Services, 
Canadian Museum of Nature, for background papers on 
Beavers. We thank the following staff of the National 
Capital Commission, Gatineau Park: Michel Viens, 
Senior Manager, Natural Resources and Land Develop- 
ment, for a permit to conduct this research in Gatineau 
Park, and Isabelle Beaudoin-Roy, Biologist, for provid- 
ing background information and reports, and, in addi- 
tion, Michel Leclair, contractor for Beaver management 
in the Park, for information on Beaver activities and 
management strategies. 


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level rise and tree mortality in a Michigan bog. Michigan 
Botanist 18: 91-98. 

Schwintzer, C. R., and G. Williams. 1974. Vegetation 
changes in a small Michigan bog from 1917 to 1972. Amer- 
ican Midland Naturalist 92: 447-459. 

Wright, J. P., C. G. Jones, and A. S. Flecker. 2002. An 
ecosystem engineer, the beaver, increases species richness 
at the landscape scale. Oecologia 132: 96-101. 


Received 28 October 2004 
Accepted 17 August 2005 


APPENDIX: VASCULAR PLANTS AND MOSSES OF THE BLACK LAKE WETLAND 


2002-2003 


Nomenclature (except for grasses) and arrangement follow Newmaster et al. (1998); nomenclature for grasses 
follows Dore and McNeill (1980). The abundance of each species is ranked on a subjective scale from rare 
through occasional, frequent and common. An asterisk (*) indicates an alien species (Gillett and White 1978). 


1. Partially Floating and Recently Beached Islands: 2002-2003 


BRYOPHYTA (Mosses) 

Sphagnum magellanicum Brid. — common 

Sphagnum teres (Schimp.) Angstr. in Hartm. — occasional 
Campylium sp.— rare 


POLYPODIOPHYTA (Ferns) 


THELY PTERIDACEAE 
Thelypteris palustris Schott — frequent 


MAGNOLIOPHYTA (Flowering Plants) 
MAGNOLIOPSIDA (Dicotyledons) 


BETULACEAE 
Alnus incana (L.) Moench ssp. rugosa (Du Roi) Clausen — 
rare, one seedling 


GUTTIFERAE 
Triadenum virginicum (L.) Raf. — occasional 


DROSERACEAE 
Drosera rotundifolia L. — rare 


VIOLACEAE 
Viola macloskeyi F. E. Lloyd ssp. pallens (Banks ex DC.) 
M. Baker — rare 


ERICACEAE 
Chamaedaphne calyculata (L.) Moench — common 


PRIMULACEAE 
Lysimachia thyrsiflora L. — rare 


LAMIACEAE 
Lycopus uniflorus Michx. — rare 


ASTERACEAE 
Bidens sp. — rare 
Eupatorium perfoliatum L. — rare 


LILIoPsIDA (Monocotyledons) 


CYPERACEAE 
Carex lasiocarpa Ehrh. — occasional to common 
Carex pseudo-cyperus L. — rare 


TYPHACEAE 
Typha latifolia L. — rare 


IRIDACEAE 
Tris versicolor L. — occasional 


ORCHIDACEAE 
Pogonia ophioglossoides (L.) Juss. — occasional in 2002; rare 
in 2003 


394 


2. Peaty Shore 2003 


MAGNOLIOPHYTA (Flowering Plants) 
MAGNOLIoPSIDA (Dicotyledons) 


NYMPHAEACEAE 
Nuphar variegata Durand in Clinton — occasional, shallow 
pool zone; remnant, non-flowering plants 


POLYGONACEAE 
*Polygonum persicaria L. — rare, upper shore zone 


GUTTIFERAE 

Hypericum mutilum L. — frequent, upper shore zone 

Triadenum virginicum (L.) Raf. — occasional, upper shore 
zone 


VIOLACEAE 
Viola macloskeyi F. E. Lloyd ssp. pallens (Banks ex DC.) 
M. Baker — occasional, upper shore zone 


ROSACEAE 

Fragaria virginiana Miller — occasional, upper shore zone 
Potentilla norvegica L. — occasional, upper shore zone 
*Potentilla recta L. — rare, upper shore zone 


ONAGRACEAE 
Epilobium ciliatum Raf. — rare, upper shore zone 
Epilobium coloratum Biehler — rare, upper shore zone 


BALSAMINACEAE 
Impatiens capensis Meerb. — frequent, upper shore zone 


APIACEAE 
Cicuta bulbifera L. — rare, upper shore zone 
Hydrocotyle americana L. — rare, upper shore zone 


LAMIACEAE 

Lycopus uniflorus Michx. — occasional, upper shore zone 

Scutellaria galericulata L. — rare, upper shore zone; common 
on unused beaver lodge 


PLANTAGINACEAE 
*Plantago major L. — occasional, upper shore zone 


SCROPHULARIACEAE 
*Euphrasia stricta D. Wolff ex Lehm. — occasional, upper 
shore zone 


CAMPANULACEAE 
Lobelia inflata L. — occasional, upper shore zone 


RUBIACEAE 
Galium tinctorium L. — occasional, upper shore zone 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


ASTERACEAE 

Bidens cernua L. — frequent, upper shore zone 

Bidens connata Muhl. ex Willd. — occasional, upper shore 
zone 


Erigeron philadelphicus L. — rare, upper shore zone 

Eupatorium maculatum L. — occasional, upper shore zone 

Eupatorium perfoliatum L. — occasional, upper shore zone 

Euthamia graminifolia (L.) Nutt.—occasional, upper shore 
zone 

Solidago canadensis L. — rare, upper shore zone 


LILiopsiDA (Monocotyledons) 


ALISMATACEAE 
Sagittaria latifolia Willd. — frequent, shallow pool zone 


POTOMAGETONACEAE 
Potamogeton epihydrus Raf. — rare, shallow pool zone 


JUNCACEAE 

Juncus brevicaudatus (Engelm.) Fern. — frequent, shallow 
pool zone 

Juncus canadensis J. Gay ex Laharpe — frequent, shallow 
pool zone 


CYPERACEAE 

Carex bebbii (L. H. Bailey) Olney ex Fern. — frequent, upper 
shore zone 

Carex pseudo-cyperus L. — rare, upper shore zone 

Dulichium arundinaceum (L.) Britton — frequent, shallow 
pool zone 

Eleocharis obtusa (Willd.) Schult. — rare, shallow pool zone 

Eleocharis sp. 

Scirpus cyperinus (L.) Kunth — rare, upper shore zone 


POACEAE 

Agrostis scabra Willd. — frequent, upper shore zone 

Torreyochloa pallida (Torr.) Church var. fernaldii (A. S. 
Hitche.) Dore — rare?, upper shore zone 


SPARGANIACEAE 
Sparganium americanum Nutt. — common, shallow pool zone 


TYPHACEAE 
Typha latifolia L. — occasional, shallow pool zone, lake edge 


IRIDACEAE 
Tris versicolor L. — occasional, upper shore zone 


Long-eared Owls, Asio otus: A Review of North American Banding 


C. STUART HOUSTON 
863 University Drive, Saskatoon, Saskatchewan S7N 0J8 Canada 


Houston, C. Stuart. 2005. Long-eared Owls, Asio otus: A review of North American banding. Canadian Field-Naturalist 
119(3): 395-402. 


In Saskatchewan, at the northern edge of the species’ range, the Long-eared Owl (Asio ofus) is an irruptive species that has 
appeared in numbers during major vole (Microtus) outbreaks in 4 of 44 years. Seven other years have had either no band- 
ings or no sightings over large areas. In a search for possible evidence of food-based nomadism, population trends, length 
and synchronicity of cycles, and longevity, I reviewed all banding in North America through 1998. Prior to banding office 
computerization (retroactive to 1955), 803 Long-eared Owls had been banded with 33 band encounters (4.1%). Between 
1955 and 1998 there were another 10 250 banded by 426 banders with 86 band encounters (0.8%). Encounter records, espe- 
cially those involving unexpected directions and distances, tend to support but do not prove food-based nomadism. Peaks of 
migration movements at different long-term stations occurred in different years. There is soft evidence of both 10- and 3-yr 
cycles. There is inconclusive evidence for a continuing population decline. The oldest banded bird in North America lived 
for 11 years, | month, but another possibly lived for 15 years, 8 months. 


Key Words: Long-eared Owl, Asio otus, nomadism, irruption, Microtus, Short-eared Owl, Asio flammeus, Northern Harrier, 


Circus cyaneus, Saskatchewan. 


In Saskatchewan, the Long-eared Owl, Asio otus, 
shows extreme variations in numbers, common only 
during obvious but unmeasured increases in vole 
(Microtus sp.) numbers. In other years, none may be 
seen across substantial areas. 

Are Long-eared Owls nomadic in North America 
as they are in Europe? (Korpimaki and Norrdahl 
1991). If so, how far do they travel in search of voles 
in high vole years? The subsequent year, when voles 
are low and the owls virtually disappear from favoured 
Saskatchewan habitat, where have the owls gone? 

The first of four long flights by banded owls in un- 
expected directions in North America was reported 1n 
1939 (Lincoln 1939); do these represent atypical flights, 
unusual dispersal, or true nomadism? If nomadic, do 
peaks occur in different years in different regions? Are 
Long-eared Owl populations cyclic, and if so, are they 
on a circa 10-yr cycle that roughly parallels Snow- 
shoe Hares (Lepus americanus) and Great Horned Owls 
(Bubo virginianus) (Houston 1987) or on a circa 4-yr 
cycle with small rodents, as occurs in Europe? Because 
this is a secretive owl, might single individuals or non- 
breeding pairs be present but overlooked during low 
years? Are North American populations of the Long- 
eared Owl steady or declining? 

I hoped North American banding data might shed 
light on some of these questions, as well as on longevity 
of this species. In particular, would banding records 
show recoveries in subsequent breeding seasons at a 
distance from the natal site? 


Methods 

The Bird Banding Laboratory of the U.S. Geological 
Survey, which maintains data for the North American 
Banding Program, provided computerized lists with 


details of all bandings and all reported band encoun- 
ters since 1955. Numbers of Long-eared Owls band- 
ed each year, 1920-1954, were obtained by perusal of 
all issues of Bird Banding Notes (Fish and Wildlife 
Service 1922-1965*), yielding 802 Long-eared Owls 
banded before computerization in 1955. Most early 
bandings were of nestlings, although a specific code 
for flightless young, age code “04, local,” was not des- 
ignated by the banding office until the August 1949 
issue of Bird Banding Notes. 

(Three 1930s bandings in Alberta were changed 
from age code “2” to the later designation of age code 
“4” in Table 2). Another 10 250 Long-eared Owls were 
banded by 426 different banders after 1955 (Table 1, 
Figure 1). Of these, 920 were of unknown age, 3499 
were locals, 2075 were immatures, 3293 were adults, 
230 were SY (second year), 231 ASY (after second 
year), one TY (third year) and | ATY (after third year). 
Of the 3499 locals banded, Idaho led with 706, fol- 
lowed by Saskatchewan (699), Alberta (361) and Mon- 
tana (317). 

Because of the intense concentration of Saskatche- 
wan banding in five different years, I paid special 
attention to the Saskatchewan subset and the rela- 
tions of their numbers to obvious vole peaks. At large 
migration stations, banding effort, year to year, of all 
ages of owls, was more consistent than elsewhere, 
including 1938 banded in Minnesota (1783 by David 
Evans at Duluth) and 1261 in Michigan (772 at 
Whitefish Point under four different permits). 


Results 
Band encounters: where and how found 

Pre-1955 banding resulted in 33 band encounters 
(4.1%). An Alberta subset of 69 nestlings and seven 


395 


396 THE CANADIAN FIELD-NATURALIST Vol. 119 


TABLE |. Long-eared Owl banding by state and province 


All-age Recoveries Banded Banded 
pre-1955 post-1955 Locals Total State or Province Main banders (>80%) of locals 
6 706 976 Idaho Marks 285; Ulmschneider 227 
| 8 698 705 Saskatchewan Houston 539; Gehlert 76 
8 l 361 399 Alberta Fyfe 160; Gehlert 19 
4 Si 864 Montana D Holt 285 
p) 284 296 Oregon PWRC #124 
] 3 DAT 301 California Bloom 175 
D 151 178 Colorado Ward 85 
l 145 158 Washington 
3 12 84 734 Wisconsin 
i=) - 90 British Columbia 
69 96 Utah 
3 14 59 1261 Michigan 
9 51 605 New Jersey 
36 65 Arizona 
3 30 46 North Dakota 
5 8 29 586 Ontario 
2 pH | 43 Pennsylvania 
11 26 1938 Minnesota 
26 pig | Nevada 
2. 3 21 519 New York 
1] 29 Iowa 
11 iS Virginia 
2 10 15 Manitoba 
10 37 Nebraska 
9 16 Kansas 
8 37 Nova Scotia 
7 50 Ohio 
a DA Quebec 
6 19 Connecticut 
2 2 4 18 Massachusetts 
3 9 Maryland 
3 8 Vermont 
2 48 Illinois 
l 6 Indiana 
l 1 New Brunswick 
0 Atlantic Ocean 
0 5 Maine 
0) 8 Missouri 
0 3 New Mexico 
0 4 Oklahoma 
0 l South Carolina 
0 8 Texas 
0 4 Wyoming 
86 3499 10250 Banded after 1955 
b)6) 802 Banded 1920-1954 
119 11052 GRAND TOTAL 34 STATES, 8 PROVINCES AND ATLANTIC OCEAN 


*Patuxent Wildlife Research Center 

Note: 57% of locals banded by 8 of 172 banding permits 

None banded in Delaware, District of Columbia, New Hampshire, North Carolina, Rhode Island, Kentucky, South Dakota, 
or in far south 


adult females banded from 1930 to 1937 by W. Ray 
Salt of Rosebud, Alberta, resulted in a remarkably high 
six recoveries (7.9%). After 1955, there were only 89 
band encounters throughout North America (0.9%). 
Of 119 band encounters from the two periods com- 
bined, 53 were found dead, 23 shot (12 pre-1955 and 
11 post-1955), 8 killed by a car or on a highway, 5 were 


injured; 18 other encounters fitted into 11 other coded 
categories, including 4 re-trapped in the same block 
of latitude and longitude in a subsequent year, 4 netted 
or re-trapped in an adjacent block, and 4 re-trapped 
in a distant ten-minute block of latitude from where 
banded. Details of the latter four travelers are: a bird 
of unknown age but probably a nestling, banded on 


2005 


Number banded per year 


TABLE 2. Long-eared Owl recoveries banded anywhere in North America. 


Band # 


000556933 
053526629 
060617751 
057690565 
0002088 16 
055674536 
060684563 
003661927 
002573140 
068604234 
003661987 
081606045 
068600215 
036714113 
040512026 
088610415 
220604277 
081611315 
081611772 
2206008 12 
081623324 
056695514 
06860021 1 
052624634 
056695534 
081698373 
220615419 
081625890 
074546503 
081693230 
038620218 
081620711 
081620719 
081641608 
068600523 
220604294 


—_ _ 
Oi CO or 
So . ©On= & 


300 
250 
200 


| 


HOUSTON: LONG-EARED OWLS 


Year 


FIGURE |. Numbers of Long-eared Owls banded each year in North America, 1955-1999. 


Distance 


(km) 


175 
225 
245 
400 
480 
590 
690 
730 


Mo 


— 


OQ WOW NK HOR RF DNAANINTONAWAOAANNWOONN KK He KN 


— 


— 


— 


Recovery Data 


Dy . Year 


a7 L932 
09) 1952. 
30. 1969 
22 1990 
IS 1933 
Ow 1957 
Zo OGY 
42 1936 
6.1936 
1980 
1934 
1O77 
1976 
1940 
1944 
1998 
1994 
1978 
1982 
1994 
198] 
1968 
Be iS 
1962 
1969 
1988 
199] 
1988 
1983 
1990 
1939 
1979 
1978 
1984 
1972 
1995 


— 
“-«* 
— 


tr bh 


\o = & WwW 
OONNOWHAANK NANA HO 


— ~] MN OO 


ON 
WwW © 


YN 
\O 


t 


NW WN 
oo WO CO 


~~ 


b 


How 


0 

3 
15 
| 
o9% 
I 
14 


45 
98 


Jes 
j= 


Nn Nn 


\O 
Seo Oia 4S ah) SS) Serre J) = S Sra S © 


TS 


State/ 
Prov. 


> 


NN DDD BD & eS Se Be Se Se Se Se KK COCOCOCCOCCOCOlUCocmclcCCcUcmcCUlcCUCCo?} 


State/ 
Prov. 


Banding Data 


Dat 


sy) 
433 
433 
435 
412 
434 
423 
424 
495 
435 
330 
465 
464 
484 
463 
433 
465 
464 
464 
43] 
464 
464 
464 
aa 
464 
464 
472 
464 
464 
472 
430 
465 
465 
465 
444 
465 


Long Mo 


1125 
875 
874 
765 
707 
791 
802 
841 
970 
785 

1170 
920 
845 

1020 

1203 
874 
920 
845 
845 
774 
845 
845 
845 
792 
845 
845 

1140 
845 
845 

1140 
892 
920 
920 
920 
875 
920 


6 
10 
10 


— 


— ea 


alla lil : | 


° 1955 Ieee 190s 19071971 1970 Taro 1963 1937 1991 1995 1999 


NM Ww 


boon NN 
NUMAN OAANWOWON NON OWANOeOC! 


—=N — N — 


i) 


_ 


397 


Year 


1931 
1959 
1969 
1989 
i927 
1958 
1965 
1932 
1936 
1978 
1934 
i973 
1976 
1937 
1943 
1989 
1993 
[977 
1981 
199] 
1979 
1966 
1975 
1954 
1968 
1988 
1990 
1985 
198] 
1989 
1939 
1978 
1978 
198] 
197] 
1993 


398 THE CANADIAN FIELD-NATURALIST Vol. 119 


TABLE 2. Long-eared Owl recoveries banded anywhere in North America (continued from previous page) 


Recovery Data Banding Data 


Distance State/ State/ 
Band # (km) Mo Dy Year How Prov. Lat Long A 5S). .Prov: Lat. Long. Mo Dye Year 
081620719 225. 10 28 1978 l WI 450 905 Z 0 MN 465 920 9 27? ISTS 
081641608 240 3 29 1984 45 WI 450 902 y 0 MN 465 920 - 10 18 1981 
068600523 355 l 24. 1972 3 Iie 413. 88 z 0 Wl 444 87> It ~ 197 
220604294 375 3 28 1995 0 MN 444 954 2 0 MN 465 920. 10 23, £983 
081604937 440 4 29. 1976 $9" MI 464 845 2 0 WI 432 S7>. 10 15., 1974 
081620400 875 f/ 12, £1978 455 "MB 5520 bol 2 0 MN 465 990) “Li 4 1976 
074575003 = 890 Ale #1992 0 IN 403 863 y) 0 ON 482 885 G 7 29 199i 
081653319) 4260 10 28 “1992; 0 NB 474 650 2 0 NJ 385 F45~ Tt 19 1988 
081620108 1305, 12 1 1978 O07 AR 352 » 935 2 0 MN 465 920 10 ioe TORS 
036714117 -—.205 8 2 py A937. | SK 502 1043 3 0 ND 484 1020 6 9 1937 
000226052 290. 11 21 1925 l IN 413 873 6) 0) MI 424 842 5 19 1925 
039669347 8330 6 28 1942 0 OH 412 814 3 0 PA 404 q15 5 16 1941 
037642104 @15, tf! 28 1938 | SD 432 ‘995 3 0) ND 484 1020 6 16 741938 
004616735 845 9 4, 1933 47 .QU 483 _770 3 0 MI 430 843 5 14 1933 
038687566 1400 12 5. 1943 0 TX 332 994 3 0 WIl 430 892 5 I5 1939 
220616638 255 9 41 1994 O MT 443 1122 4 0 MT 463 1140 5 7 1991 
057692535 280 10 8 1968 O NY 424 £734 4 0) NJ 402 750 5. 25. 1968 
057689020 435 6 28 1969 4 MB: 505.1005 4 0 SK 520091 10653 6 15 .. 1967 
057688448 715 ys 10 1967 3 MT 454 1083 4 0 SK 520 1064 6 9 1966 
057692533 995 10 4 1969 | NB 473 6/72 4 0 NJ ~ 402 745 > (20 1968 
004629600 1135 z 2 1935 l UT 410 1115 4 0 AB Sit) 1135 if 4. +933 
057688480 1415 4 9 1969 0 TA 435. +935 4 0 SK: 5207 1070 6 , 20, "1866 
004629579 1440 3 3 1934 98 NE 425 981 4 0 AB Sik 1125 fl 4 1933 
050643309 1605 4 1 1961 | AB. 524 1133 4 0 CO 393 1045 6 2 1959 
003664770 1660 10 21 1933 98 MN 442 930 4 0 AB’ Sil » 1225 6. 25, 1935 
057689017 2365 12 99 1972 0 MS. 3241. 9035 4 0 SK 520 1064 6 I> 1967 
057695906 2385 = 11 3. 1957 | NL 263 1001 4 0 MI 433 840 5) lL? 1957 
056681737 2825 8 64 1972 98 NL 254 1001 4 0 Sk 510 ‘1022 6 20° a7 
057612231 3710 | 25 1970 | DF 191 990 4 0 SK 520 1064 6 13 1969 
052622635 3865 99 99 1960 0 OA 163 964 4 0 SK’ 510° 1020 6 15 1960 
081611823 465 4 41 1997 0 Mi 421 «33 5 4 NY ~ 431 774 4 20 1996 
074597414 645 7 5 1999 0 ON 463 = 842 6 0 NY. 435i 774 4 5 1989 


In column 6, how found codes are: 0, found dead; 1, shot; 2, injured; 3, starved; 4, caught in trap; 14, struck by vehicle; 15, 
killed by weather; 45, killed on highway; 47, band removed; 50, skeleton only; 57, entangled in fence; 89, trapped and 
released in different 10-minute block; 98, band only without information. 


In column F. all bird codes indicate death, except for 89* 


In columns 7 and 12, standard postal abbreviations for states and provinces are used. 


Additional abbreviations for Mexican states in column 7 are: 


DF — Federal District; GU — Guanajuoto; NL — Nuevo Leon; OA — Oaxaca; PU — Puebla; QU — Queretaro 


17 April 1932 by P. F. English in Michigan, was re- 
trapped during mid-February 1936 in Virginia, 730 km 
distant; an immature banded in Wisconsin by C. R. Sin- 
delar on 15 October 1974 was re-trapped on 29 April 
1976 in Michigan, 440 km distant; an owl of unknown 
age, banded at Nature Dunes Nature Center, Wisconsin, 
9 November 1981, was re-trapped on 30 November 
1982, 130 km distant in Wisconsin; a nestling banded 
at Rosebud, Alberta, by Ray Salt on 25 June 1931, was 
re-trapped 175 km north and west near Killam, Alberta, 
in mid-May 1932. All but the Alberta ow! were prob- 
ably in migration when banded. 

The map (Figure 2) and Table 2 together depict the 
movements of 64 owls that traveled more than 100 km. 


Fifteen of the 64 were banded as nestlings, and trav- 
eled greater distances, an average of 1672 km, than 
owls banded at other ages. One wandered north 280 km 
from New Jersey, in the months after it was banded. 
Another two were shot after “wrong-way-Corrigan” 
northward movements of 1605 and 995 km, respective- 
ly: from near Denver, Colorado, banded by J. A. Neff 
on 2 June 1959, to Ponoka, Alberta on | April 1961, 
and from near Stockton, New Jersey, banded by O. A. 
Heck on 20 May 1968, to Saint Quentin, New Bruns- 
wick on 4 October 1969. The other 12 nestlings trav- 
eled in a southerly direction, 4 reaching Mexico along 
with 3 of other ages. Once could speculate that the 
seven recoveries in Mexico might represent “leap- 


2005 


HOUSTON: LONG-EARED OWLS 


Ete 


LEOW 


e@ Banding locations 
* Recovery locations 


FIGURE 2. Map of Long-eared Owl movements in North America, 1925-1999. Circles represent place of banding. Stars rep- 


resent place of recovery. Map by K. M. Meeres. 


frogging” of northern owls to more southern winter- 
ing grounds. 

There are four other examples of dispersal in an 
unusual direction and distance. The first two, presumed 
to be young in their first year of life, were each recov- 
ered in the year of banding. Lincoln (1939: 128) listed 
a Long-eared Owl of unknown age banded at Escon- 
dido, California, 22 April 1934, and shot at Corbeil, 
Ontario, 9 October 1934, a distance of 3135 km north- 
east. Considering banding of all species, Lincoln com- 
mented “This is one of the most unusual records thus 
far obtained.” Lincoln also listed a juvenile Long-eared 
Ow/l that had traveled northeast from St. John, Michi- 
gan, to Abitibi, Quebec, between 14 May and 4 Sep- 
tember 1933, a distance of 845 km. In each of these 
instances it is not known whether the northward move- 
ment was achieved in the first or second year of life, 
nor whether either represents migratory movement or 
dispersal to a very different breeding location. 

Four distant recoveries during or immediately after 
the breeding season might have been construed as evi- 
dence in favor of nomadism, had they not been banded 
during migration. 

Two were adults banded as April migrants at Brad- 
dock Bay, New York; one flew 465 km to the west to 


Michigan in mid-April of the subsequent year and the 
other was found dead 645 km northwest in Ontario 
on 5 July, 10 years after it had been netted as an SY 
bird by subpermittee Frank Nicoletti (see below under 
Longevity). Two others, banded as late migrants at 
Whitefish Point, Michigan (on 22 April and 14 May), 
were recovered two years later; one went northeast 
480 km and was shot in mid-June in western Quebec; 
the other went southeast 470 km and was found dead 
on 30 July in Ontario. 


Longevity 

After deleting ten birds inadmissible because they 
were alive when recaptured (most of them taken in a 
net at a banding station and released), two bands found 
on a skeleton, and two reported as “band only,’ 105 
records remained that were acceptable for a life table 
(Table 2). As with other raptor species, nearly two- 
thirds of the band encounters, 67 of 105, occurred 
before the birds were a year old, with 17 in the sec- 
ond year (Table 3). 

The Long-eared Owl accepted by the banding office 
as their oldest to date was banded as a second-year bird 
at Braddock Bay, New York, by Frank Nicoletti on 5 
April 1989, and found dead in Ontario on 5 July 1999, 
as already mentioned. At least a year old when banded, 


400 


assuming hatching near the first of June, it was at 
least 11 years, | month old, recognized as the longevi- 
ty record for this species (Klimkiewicz 2005*). A 
potentially older Long-eared Owl is also in the band- 
ing records. Number 816-06355, banded as a nestling 
near Lincoln, Nebraska, on 24 May 1978 by Ross 
Lock, was reported as killed by a hawk or owl on an 
unknown date in February 1994, in the same ten- 
minute block of latitude and longitude in which it had 
been banded. Long-eared Owls regularly winter that 
far north. The unusual and precise mode of death tends 
to suggest that the band had not been lying in a desk 
for months or years before it was reported. Had this 
owl indeed died in February 1994, it would be the 
oldest Long-eared Ow] in North America, at 15 years, 
8 or 9 months. Older birds are expected in future, since 
in Europe, the longevity record for this species 1s 27 
years, 9 months (Rydzewski, cited in Marks et al. 
1994). 


Relation to vole numbers in Saskatchewan 

In Saskatchewan, between 1969 and 1997, only in 
three years (1970, 1982, and 1986) were no Long- 
eared Owl nests detected anywhere in extensive 
travels throughout west-central Saskatchewan (CSH, 
personal observation). Also in 1982, and in three 
other years (1976, 1979, and 1981), no birder in the 
11 012 km? Saskatoon area reported seeing a single 
Long-eared Owl at any time (Leighton et al. 2002: 
182-183). 

The three peak vole years (1960, 1969 and 1997) 
made a striking contrast. Each occurred in a spring 
following the rare occurrence of grain crops that were 
incompletely harvested before snowfall. Some grain 
crops lay in the swath unharvested all winter, and voles 
multiplied in the grain beneath the snow. Greatly in- 
creased numbers of breeding Long-eared Owls ap- 
peared in each of the following springs, together with 
increased numbers of Short-eared Owls (Asio flam- 
meus) and Northern Harriers (Circus cyaneus). In 
1960, there was a striking increase in the numbers of 
breeding pairs of Long-eared Owls (73 nestlings band- 
ed), Short-eared Owls (68 nestlings) and Northern 
Harriers (25 nestlings). In 1960, productivity of the 
Great Horned Ow! peaked in concert that year with the 
ten-year peak of the Snowshoe Hare (Houston 1960). 

In the spring of 1969, banders capitalized on a 
recurrence of high vole numbers to band 195 Long- 
eared Owl nestlings, 104 Short-eared Owl nestlings 
and 202 Northern Harrier nestlings (Houston 1997). 
Long-eared Owl sightings by members of the Saska- 
toon Nature Society also peaked dramatically in 1969, 
reaching levels four to ten times higher than any year 
before or since (Figure 1 in Houston 1997). 

Although not as widely spread as in 1959 and 1968, 
occasional fields of grain lay unharvested at first snow- 
fall in the fall of 1996. In the subsequent spring, 1997, 
voles were common, based on direct observation and 
anecdotal reports. Some days on Highway 6 south of 


THE CANADIAN FIELD-NATURALIST 


Vol Tig 


TABLE 3. Long-eared Owl life table. 


8-9 
9-10 
10-11 
15-16 
Total 


“ig 
(o°e) 
S) 
A=—_— ON OK NN F CO 


119 band encounters 10 inadmissible, because alive when 
recaptured; 4 excluded, because skeleton (code how found, 
50) or 98 (band only), after one year. 105 records available 
for life table. 


Regina, voles crossed the highway in such numbers 
that vehicles left a patchwork of dead voles on the 
asphalt surface. The three mouse-eating raptors that 
year showed only modest increases in banding of 
nestling Long-eared Owls (18), Short-eared Owls (15), 
and Northern Harriers (23). Fifteen of these Long- 
eared Owls were on the Marten J. Stoffel (MJS) dairy 
farm raptor study area immediately north of Saska- 
toon, a rectangle 8 km east to west and 13 km north 
to south. Here MJS had found no nesting Long-eared 
Owls in 1998, one pair with four young in 1999, and 
an unprecedented density of 36 pairs of breeding Long- 
eared Owls, one pair per 2.9 km?, in 2000 (Stoffel 
AUOT): 


Are irruptions synchronous ? 

Unlike Great Horned Owls, whose 10-year cycle 
tends to be roughly synchronous across much of North 
America (Houston 1987), peak years for Long-eared 
Owls appeared to vary widely between localities. At 
Duluth, D. L. Evans banded 172 Long-eareds in 1978 
and 163 in 1981, and mist-netted more than 100 dur- 
ing migration in three other years, 1980, 1986 and 
1990. At Whitefish Point, Michigan, the overwhelm- 
ing peak year was 1981. At Cedar Grove, Wisconsin, 
the peak year was 1982, whereas Erdman’s station near 
Oconto, Wisconsin, had peak numbers in 1977. At 
Camden, New Jersey, the peak year was 1988, and at 
Braddock Bay, New York, the peak years were 1993 
and 1994. 


How pronounced are cycles? 

A casual glance at Figure | offers only weak sup- 
port for a ten-year cycle. However, analysis by James 
R. Duncan (Manitoba Natural Resources, personal 
communication) showed 14 minor peaks and hence 
13 between-peak-periods (BPP). He found four “2-yr 
BPPs,” 5 “3-yr BPPs,” and four “4-yr BPPs.” Thus, 
the mean and coincidentally, the mode BPP was 3 


2005 


years. Further, Evans’ Figure 6, in Marks et al. 1994, 
shows a four-year cycle of incomplete molt. 

Numbers banded at migration stations, where annu- 
al effort is more consistent than in itinerant travels to 
locate nests containing young, showed a continuing 
decline in Long-eared Owls captured. Apart from band- 
ing, there is also suggestive evidence for declines in 
California, Indiana, Maryland, Minnesota, New Jersey, 
Pennsylvania, and South Dakota, but not for western 
Montana (Holt 1997). This suggestion of a gradual 
decline, even better documented for Short-eared Owls 
from Breeding Bird Survey data, is somewhat at odds 
with the locally high numbers seen in Saskatchewan 
in 2000. 


Discussion 

Long-eared Owl numbers in Saskatchewan fluctu- 
ate in relation to vole numbers; during seven years 
these owls virtually disappeared from major portions 
of the province. This was not the case in western Mon- 
tana or southwestern Idaho, where they were found 
year-round, including winter, every year, especially in 
sheltered valleys. On Denver Holt’s study area north 
of Missoula, Long-eared Owls were present and band- 
ed every year since 1987, although there was some 
fluctuation in numbers and in 2 of 15 years (1993 and 
1995) he banded no nestlings. In southwestern Idaho, 
Helen Ulmschneider (1993, 1994) and John Doremus 
banded young in successful nests for 12 consecutive 
years, 1988 through 1999. 

What might explain the differences between Saska- 
toon, Saskatchewan (52°N), where Long-eared Owls 
do not winter, and Holt’s study area (47°N) in Mon- 
tana? Because both Saskatchewan and Montana have 
adequate numbers of corvid nests, nest sites are not a 
limiting factor. Even in extreme southwestern Saskat- 
chewan where there are sheltered wooded valleys, 
Long-eared Owls rarely remain through Christmas 
week. In contrast, large winter roosts are encountered 
in Idaho and Montana and at least some owls from 
the winter roosts remain to nest (Ulmschneider 1993, 
1994; D. W. Holt, personal communication). Owls 
occupy pastoral inter-mountain valleys in Montana 
whereas the Saskatoon area is largely a cultivated, 
wind-swept plain with pastures for dairy cattle and 
scattered copses of aspen (Populus sp.). 

Because vole cycles are known to be more pro- 
nounced in northern Fennoscandia than farther south 
in Europe (Korpimaki and Norrdahl 1991), and Long- 
eared Owls winter farther north in Finland in years of 
vole abundance (Korpimiaki 1994), it is possible that 
vole fluctuations are more extreme in southern Sas- 
katchewan than in Montana, although I know of no 
Saskatchewan mammal trapping data to support such 
an hypothesis. In Finland, there is no time lag between 
population fluctuations of vole-eating raptors and their 
prey (Korpimaki 1994). 


HOUSTON: LONG-EARED OWLS 


401 


When Long-eared Owl numbers peaked at or near 
the top of the ten-year Snowshoe Hare cycle in 1960 
and 1969, Great Horned Owls had ample hares to feed 
their young. In such years, Great Horned Owls had less 
need to take voles, and perhaps competed less often 
with Long-eared Owls for that prey base. 

What have I learned from this review? Clearly, 
Long-eared Owls and Short-eared Owls increase when 
voles increase, and thus are owl equivalents of the 
Northern Harrier, which Fran Hamerstrom (1986) 
termed “the hawk that is ruled by a mouse.” Apart from 
its clear relationship to vole numbers, other answers 
are as yet incomplete: 

1. Are Long-eared Owls nomadic? Movements of 
individual banded Long-eared Owls offer tantaliz- 
ing clues rather than strong evidence in support of 
nomadism, at least in the northern portions of the 
range. Full proof of nomadism requires that marked 
birds that bred in one area be sighted or captured 
while breeding in another area at a considerable 
distance. The Northern Saw-whet Owl, for exam- 
ple, appears to settle to breed in areas of high food 
abundance that it encounters during the nonbreed- 
ing season (Marks and Doremus 2000). 

2. Are peaks synchronized? No. Numbers of Long- 
eared Owls peak in different years at different migra- 
tion stations. 

3. Length of cycle? There is “soft” evidence for both 
three- and ten-year cycles. 

4. Is there an overall, continuing population decline? 
Not proven. Evidence from migration stations, 
especially at Duluth, 1976-1993 (Evans’ figure 5 
in Marks et al. 1994), suggests yes; Evans (person- 
al communication) informs me that the average of 
90 per year banded at his station, 1974-1993, 
dropped to 31 for 1991-2003. The evident concen- 
trations that appeared in Saskatchewan in three 
different years do not exclude a long-term, general 
downward trend in population. 


As Marks et al. said in 1994, similar questions “can 
be answered only by intensive banding and recapture 
efforts over many years.” Here is an opportunity for 
all banders to band both nestlings and adults whenever 
possible. Application of readily visible wing tags or 
satellite radiotelemetry, should funds and miniatur- 
ization permit, might prove invaluable. All of us have 
a great deal yet to learn about this enigmatic and per- 
plexing species. 


Acknowledgments 

I thank Gary Bortolotti, Elizabeth Brooks, Tom 
Carpenter, Jim Duncan, Dave Evans, Denver Holt, 
Jeff Marks, Bob Nero, Frank Roy, Marten J. Stoffel, 
and Dan Zazelenchuk for helpful comments. Eliza- 
beth Brooks, David Evans, and Denver Holt gave 
permission to use their banding data, as did Michael 


402 


Kochert on behalf of his subpermittees, Jeff Marks, 
Helen Ulmschneider, and John Doremus. Mary Gus- 
tafson provided the number of individuals banded by 
W. Ray Salt in Alberta in the 1930s. James R. Dun- 
can analysed the data for determination of cycle length. 
Kathy Meeres prepared the map. Ross A. Lock and his 
successor, John Dinan, at Nebraska Game and Parks, 
kindly followed up on the 15-year-old recovery record. 


Documents Cited (marked * in text) 

Fish & Wildlife Service. 1922-1965. Bird Banding Notes, 
volumes 1-5 (mimeographed). Bureau of Biological Survey, 
Washington, District of Columbia and Fish and Wildlife 
Service, Patuxent, Maryland. 

Klimkiewicz, M. K. 2002. Longevity records of North Amer- 
ican birds. Version 2002.1. Patuxent Wildlife Research 
Center, Bird Banding Laboratory, Laurel, Maryland. http:// 
www.pwrc.usgs.gov/bbI/homepage/long vrec.htm. 


Literature Cited 

Hamerstrom, F. 1986. Harrier: Hawk of the marshes. Smith- 
sonian Press, Washington, District of Columbia. 

Holt, D. W. 1997. The Long-eared Owl (Asio otus) and forest 
management: a review of the literature. Journal of Raptor 
Research 31: 167-174. 

Houston, C. S. 1960. 1960, the year of the owls. Blue Jay 17: 
105-110. 

Houston, C. S. 1987. Nearly synchronous cycles of the Great 
Horned Owl and Snowshoe Hare in Saskatchewan. Jn 
Biology and conservation of northern forest owls. Edited 
by R. W. Nero, R. J. Clark, R. J. Knapton, and R. H. Hamre. 
General Technical Report RM-142. Fort Collins, Colorado: 
U.S. Department of Agriculture, Forest Service, Rocky 
Mountain Forest and Range Experiment Station. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Houston, C. S. 1997. Banding of Asio owls in south-central 
Saskatchewan. Jn Biology and conservation of owls of the 
northern hemisphere. Edited by J. R. Duncan, D. H. John- 
son, and T. H. Nicholls. General Technical Report NC- 
190, U.S. Department of Agriculture, Forest Service, North 
Central Research Station, St. Paul, Minnesota. 

Korpimaki, E. 1994. Rapid or delayed tracking of multi- 
annual vole cycles by avian predators? Journal of Animal 
Ecology 63: 619-628. 

Korpimaki, E., and K. Norrdahl. 1991. Numerical and func- 
tional responses of kestrels, short-eared owls, and long- 
eared owls to vole densities. Ecology 72: 814-826. 

Leighton, A. L., J. Hay, C. S. Houston, J. F. Roy, and S. 
Shadick. 2002. Birds of the Saskatoon Area. Saskatche- 
wan Natural History Society Special Publication 23, Regi- 
na, Saskatchewan. 

Lincoln, F. C. 1939. The migration of American birds. 
Doubleday Doran, New York. 

Marks, J. S., and J. H. Doremus. 2000. Are Northern Saw- 
whet Owls nomadic? Journal of Raptor Research 34: 299- 
304. 

Marks, J. S., D. L. Evans, and D. W. Holt. 1994. Long-eared 
Owl (Asio otus). In The birds of North America, 133 (24 
pages). Edited by A. Poole and F. Gill. The Academy of 
Natural Sciences, Philadelphia, Pennsylvania, and The 
American Ornithologists’ Union, Washington, District of 
Columbia. 

Stoffel, M. J. 2001. Long-eared Owl abundance near Saska- 
toon in 2000. Blue Jay 59: 129-133. 

Ulmschneider, H. M. 1993, 1994. Wintering and nesting 
site use by Long-eared Owls in the Snake River Birds of 
Prey Area. Snake River Birds of Prey Area 1993 Annual 
Report: 318-323; 1994 Annual Report: 292-295. 


Received 17 December 2004 
Accepted | September 2005 


Cyclopyxis acmodonta n. sp. and Arcella formosa n. sp.: Two New 
Species of Testate Rhizopods (Arcellinida, Protozoa) from Remnant 
Wetlands in Ontario, Canada 


KENNETH H. NICHOLLS 


S-15 Concession 1, RR # 1 Sunderland, Ontario LOC 1HO Canada; e-mail: khnicholls @interhop.net 


Nicholls, Kenneth H. 2005. Cyclopyxis acmodonta n. sp. and Arcella formosa n. sp.: two new species of testate rhizopods 
(Arcellinida, Protozoa) from remnant wetlands in Ontario, Canada. Canadian Field-Naturalist 1 19(3): 403-411. 


Species of the testate rhizopod genera Arcella and Cyclopyxis are found predominantly in the benthos of shallow ponds, for- 
est pools, marshes and other freshwater habitats. Their taxonomy is based on the morphology of their tests — the shell-like 
structure that houses the living single-celled amoeboid protoplast. This paper describes a new species of Cyclopyxis discovered 
in two coniferous forest bogs, and a new species of Arcella from a Typha-dominated marsh on the north shore of Lake 
Ontario, Canada. Cyclopyxis acmodonta Nicholls n. sp. differs from its closest relative C. stellata (Wailes) Deflandre 1929 in its 
much smaller size and its irregularly shaped pseudostomal aperture, which is embellished with numerous, tiny, sharp-pointed tooth- 
like quartz granules around the interior margin. Although the saucer-like indentations over the dorsal surface of the test of A. 
formosa Nicholls, Meisterfeld & T6r6k n. sp. resemble similar structures found in several other Arcella species, the large size 
of A. formosa (165-235 Lm in diameter) and its low profile (low height-to-diameter ratio) are the main features distinguishing 


this new species from other Arcella species. 


Key Words: Arcella, Arcellaceans, Arcellidae, amoebae, Cyclopyxis, Testacealobosia, Rhizopoda, Trigonopyxidae. 


The species-level taxonomy of most testate rhi- 
zopods has been based on the size and shape of their 
shells or tests — the structure that “houses” the amoe- 
boid protoplast. Species of the testate amoebae gen- 
era Arcella and Cyclopyxis occupy similar habitats 
(generally shallow benthic freshwaters, bogs, forest 
pools and damp mosses) and possess certain superfi- 
cial similarities in their test structure relating to the 
usual dome-shaped radial symmetry, including a cen- 
trally located pseudostomal aperture. Test composition 
in these two genera is, however, very different and this 
fact has been the basis for their separate placement in 
two distinct families. Tests of Arcella (family Arcelli- 
dae) are of a rigid and transparent organic material, 
while those of Cyclopyxis (family Trigonopyxidae) are 
composed of mineral particles embedded in an organic 
cement (Meisterfeld 2002). There are presently about 
120 and 80 species and subspecies of Arcella and 
Cyclopyxis, respectively. 

Deflandre (1928) provided a monographic treatment 
of the genus Arcella that was followed by a similar com- 
pilation for the genus Centropyxis (Deflandre 1929). 
In his treatment of the genus Centropyxis, Deflandre 
(1929) erected the subgenus Cyclopyxis to include those 
Centropyxis-like tests with radial symmetry (Centropy- 
xis tests are bilaterally symmetric). Virtually all authors 
since then have considered Cyclopyxis an autonomous 
genus quite distinct from Centropyxis. Arcella and 
Cyclopyxis taxonomy has been summarily updated by 
Decloitre (1976, 1977, 1979, 1982 and 1986). Unfortu- 
nately, many taxa, especially at the subspecies level, 
were originally poorly described on the basis of very 


few specimens, so little is known of the range of form 
variation and how this relates to the morphology of 
similar but differently named taxa. Based on a detailed 
study of form variation of tests, Foissner and Korganova 
(1995) suggested that nine species and subspecies of 
Cyclopyxis might be reduced to two valid species. Since 
Decloitre’s latest taxonomic update (Decloitre 1986), 
descriptions of new taxa of Arcella and Cyclopyxis have 
been few (e.g., Chardez et al. 1987; Torres and Jebram 
1993; Balik 1995). 

The purpose of this paper is to describe one new 
species in each of the genera Arcella and Cyclopyxis 
discovered recently in wetlands in Ontario, Canada. 
Features of test morphology of both species are high- 
ly distinctive and not easily confused with previously 
known taxa. Detailed statistics on test dimensions, as 
well as light microscopic digital images and draw- 
ings, are included. 


Methods 
Sampling locations and collection methods 
Cranberry Marsh (43°50'38"N; 78°57'0"W) is a 
16-ha cattail (7ypha)-dominated wetland within the 
Town of Whitby, Ontario (Figure |). There are no per- 
manent surface inflows or outflow from the marsh, al- 
though some water exchange with Lake Ontario is pos- 
sible during storm surges across a low berm separating 
the marsh from the gravel/cobble beach of Lake On- 
tario. Cranberry Marsh is a provincially significant wet- 
land managed by the Central Lake Ontario Conserva- 
tion Authority and is an important nesting/feeding/rest- 
ing area for resident and migrating songbirds, waterfowl 


403 


404 


79° 30' 79°00' W 


Foi! 


FIGURE 1. Map showing three testate rhizopod collection 
locations between Lake Ontario and Lake Simcoe in 
southern Ontario. Latitude and Longitude for Cran- 
berry Marsh, and Sites A and B are as follows: 
43°50'38"N, 78°57'30"W; 44°08'27"N, 79°05'12"W; 
44°12'33"N, 79°04'40"W, respectively. 


and shorebirds. Samples of water and bottom sediment 
were collected 7 August 2004 by submerging a wide- 
mouthed 500 mL bottle at the south end of Cranberry 
Marsh where water depth was about 0.5 m. 

The Beaver River originates in a forest bog and 
marsh complex approximately 5 km southeast of the 
town of Uxbridge, Ontario (Figure 1). With a total 
length of approximately 50 km, a watershed area of 
3.2 x 10° m? and a total annual discharge that aver- 
aged 8.3 x 10’ m® year! for the 1990s (Scott et al. 
2001*), the Beaver River is one of the largest rivers 
flowing into Lake Simcoe. Sampling for testate rhi- 
zopods was in moss pools near the upper Beaver River 
(Sites A and B in Figure 1) in areas of forest bog 
dominated by White Spruce (Picea glauca (Moench) 
Vos, Northern White Cedar (Thuja occidentalis L., 
Tamarack (Larix laricina (Du Roi) Koch), Sphagnum 
spp, Hylocomium splendens, and other forest mosses. 
Samples containing testate rhizopods consisted of ap- 
proximately 10 cc of wet moss and 500 mL of water 
mixed with bottom sediment from small forest pools 
of 20 cm maximum depth. 


Laboratory methods 

Samples were examined shortly after collection in 
their living state with an inverted microscope. Other 
portions of the samples, comprising about | cc of 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


sediment and 20 cc of water, were fixed with formalin 
to achieve a final concentration of about 4% formalde- 
hyde. Testate rhizopods of interest for measuring, image 
capture or isolation for transfer to a separate perma- 
nent preparation were isolated from surrounding debris 
by manipulation at low power (10x objective) with a 
single hair brush. A single specimen of each of the 
two new species described here was selected to serve 
as the type specimen for museum archival and were 
transferred with a micro-pipette to a Number | cover 
glass for drying and subsequent mounting on a slide 
with Canada Balsam. All measurements were made at 
a magnification of 600x (40x objective, 1.5x micro- 
scope head, 10x eyepiece). Optimal orientation for 
measurement was achieved by manual manipulation 
of isolated specimens with a single hair brush. Des- 
criptive statistics on measurements were run in CoStat 
(CoHort Software 1995*). Images were captured with 
a 3.4 megapixel digital camera and assembled onto 
plates for publication using Adobe Photoshop 5.0. 


Results 

Cyclopyxis acmodonta n. sp. 

Phylum Rhizopoda Class, Lobosea; Order Arcel- 
linida; Family Trigonopyxidae Loeblich and Tap- 
pan, 1964. 

Diagnosis: Test constructed of agglutinated micro- 
scopic quartz granules and in side view with a flat- 
tened ventral surface and domed dorsal surface. Ven- 
tral surface with a centrally located and irregularly- 
shaped oral aperture (pseudostome) about the test 
diameter and composed of from three to seven lobes 
or indentations. Margin of the pseudostome orna- 
mented with many sharp-pointed tooth-like granules. 
Test diameter, 188-298 tm; test height, 150-217 um; 
widest opening in pseudostome (between distal 
lobes), 75-143 Um. Pseudostome only slightly invagi- 
nated relative to the surrounding ventral surface of 
the test. 

Etymology: The specific epithet (““acmodonta’’) refers 
to the small sharp-pointed “teeth” that embellish the 
interior margin of the pseudostome [acmodonta 
(Latin, fem. adj.) = sharp-toothed]. 

Type specimen: The type specimen was mounted in 
Canada Balsam on a glass slide and was deposited 
with the Invertebrate Zoology Division, Canadian 
Museum of Nature, Catalogue Number CMNI 2005- 
0004. 

Holotype material: A formalin-preserved aqueous 
sample has been retained by the author under sam- 
ple No. V-1892, collected 4 January 2004. 

Type locality: Forest bog (Sphagnum moss dominat- 
ed small pool) near the upper Beaver River, Ontario 
Canada (44°08'27"N, 79°05'12"W). Specimens were 
also collected on 16 October 2001 from a moss pool 
in a similar forest habitat (Site B in Figure 1; 
44°12'33"N, 79°04'40"W), approximately 8 km N 
of the type locality. 


2005 NICHOLLS: TWO NEW SPECIES OF TESTATE RHIZOPODS 405 


TABLE |. Sizes of test variables in Arcella formosa and Cyclopyxis acmodonta. n = 14 for all A. formosa variables; n = 18 
for C. acmodonta except n= 14 for ap and D/ap. Definitions of D, H and ap are as depicted in Figures 2a, 2b and 4b, respec- 
tively. The pseudostomal “ap” metric for C. acemodonta was represented by the longest straight line joining the most remote 
lobes of the aperture. For those A. formosa specimens for which the aperture was slightly elliptical rather than circular, the 


longest distance through the centre was measured. 


Arcella formosa diameter (D) height (H) aperture (ap) D/H D/ap 
median 192 85 58.5 25 52 
minimum 165 56 46 1.9 2D 
maximum 235 100 85 3:2 3.6 
mean 199.1] 84.9 64.6 2.4 a 
SD pA Be 12.4 15.6 0.4 0.4 
coef. var. (%) 13.9 14.9 24.5 16.2 ie! 

Cyclopyxis acmodonta diameter (D) height (H) aperture (ap) D/H D/ap 
median 260 195 118 1.4 O49) 
minimum 188 150 75 1.2 1.9 
maximum 298 pANG| 143 L5 2D 
mean 249.9 185.7 116.1 13 Or 
SD 32] ZA a ia 0.1 0.2 
coef. var. (%) 13.9 14.9 24.5 16.2 11.4 


FIGURE 2. Cyclopyxis acmodonta tests. a. Drawing of the ventral view showing the irregularly lobed and centrally located 
pseudostomal aperture (D = test diameter). b. Drawing of the lateral view; the flat side is the ventral surface (H = 
test height). ¢. Image of the type specimen archived with the Canadian Museum of Nature (Catalogue Number 
CMNI 2005-0004). The scale bar in Figure 2c applies to Figures a-g. d. Lateral view showing agglutinated quartz 
granules on the test surface. e-g. Three different specimens in lateral view showing variation in profile shape. h. 
Details of test surface showing incorporated diatom frustules (arrows). i. interior margin of one lobe of a pseudos- 
tomal aperture showing three tooth- like granules (arrows). 


406 


In lateral view, the shape of the test of Cyclopyxis 
acmodonta was quite distinctive, although there was 
some variation in the shape of the domed dorsal sur- 
face, ranging from those with a smooth rounded form 
to those with a slightly more conical shape (Figures 
2b,d-g). Certainly, the term hemispherical, which has 
been applied to several other species of Cyclopyxis, 
cannot be applied here because of C. acmodonta’s 
more exclusive shape. In a hemisphere, the diameter- 
to-height ratio is 2; in C. acmodonta, the test diameter- 
to-height ratio ranged from 1.2 to 1.5 with a median 
of 1.4 (Table 1). 

The most distinctive feature of this species is the 
pseudostome with its highly variable and irregularly 
lobed margin (Figures 2a,c, 3a-i). Higher magnification 
revealed the many tiny sharp-pointed angular quartz 
grains attached to the inner margin of the pseudostome 
rim (Figure 21), but these were also apparent in some 
specimens at lower magnification (e.g. Figures 3e,h). 
The number of apertural lobes ranged from three to 
seven (nine, if smaller subdivisions of lobes were in- 
cluded). The pseudostome was only slightly invaginat- 
ed (<1/5 test height) relative to the surrounding ventral 
surface of the test. 

In some specimens there was appeared to be a thin 
membrane-like cover over the aperture in which were 
embedded a few thin quartz particles and diatoms 
(Figure 3g). This structure may be an early component 
of cyst formation, whereby the aperture becomes more 
densely plugged at later stages of encystment. Many 
specimens were observed with the apertural rim only 
faintly visible owing to the large accumulation of test- 
like material over the pseudostome. Presumably, these 
represented a later stage of encystment. In these speci- 
mens too, the internal protoplast was dense and darkly 
coloured. 

All tests examined were completely covered in high- 
ly refractive microscopic quartzite particles. There did 
not appear to be any distributional patterns in the sizes 
of these particles, suggesting that during test construc- 
tion, the organism does not discriminate among particle 
sizes for specific regions of the test. Larger particles 
appeared to be randomly dispersed and interspersed 
with smaller particles over the whole of the test. Rarely 
were particles larger than 50 um diameter found in tests 
of this species. Intact and broken diatom frustules were 
sometimes encountered (Figure 2h), again, with no 
apparent preference for either dorsal, lateral or ventral 
surfaces of the test. 


THE CANADIAN FIELD-NATURALIST 


VoL. 119 


Arcella formosa Nicholls, Meisterfeld & Tér6ék n. 

sp.! 

Phylum Rhizopoda; Class Lobosea; Order Arcellinida; 
Family Arcellidae Ehrenberg, 1830 emend. Deflan- 
dre, 1953: 

Diagnosis: Test nearly colourless to dark rusty-brown 
in colour, constructed of circular-to-elliptical areoles, 
3.0-4.5 x 2.5-3.5 um. In ventral and dorsal views 
the test is generally circular in outline but with a 
wavy or “lumpy” margin 165-235 tm in diameter. 
The pseudostomal aperture consists of a thickened 
rim, 46-85 tum in diameter. The pseudostome is in- 
vaginated relative to the surrounding ventral surface 
of the test by a distance of about /% the test height. 
The internal rim of the pseudostome is recurved 
forming a short (3-4 um) buccal tube. In lateral view, 
the test consists of a low, dome-shaped structure 
with multiple depressions that impart a wavy-edged 
appearance to the margin of the test. Test height is 
56-100 Lum. 

Etymology: The specific epithet (“formosa’) refers to 
the attractive smooth curves created by the depres- 
sions in the dorsal surface of the test [formosa (Latin, 
fem. adj.) = beautiful]. 

Type specimen: The type specimen was mounted in 
Canada Balsam on a glass slide and was deposited 
with the Invertebrate Zoology Division, Canadian 
Museum of Nature, Catalogue Number CMNI 2005- 
0003. 

Holotype material: A formalin-preserved aqueous 
sample has been retained by the author under sam- 
ple Number V-1924, collected 7 August 2004. 

Type locality: South end of Cranberry Marsh, Town of 
Whitby, Ontario, Canada (43°50'38"N, 78°57'30"W). 
The large size (median test diameter of 192 um; 

Table 1) and the “lumpy” appearance of the test mar- 

gins, in both ventral and lateral views (Figures 4a, b), 

are distinctive features of this species. There was con- 

siderable variation in the colour and degree of develop- 
ment of the depressions on the dorsal surface of this 
species. In the lighter coloured specimens (pale yel- 
lowish-grey), the dorsal depressions were less well 
developed (Figures 4c-f) than in the darker (brownish 
orange) coloured specimens, where the depressions and 
the thickened rims delineating their boundaries were 
very well defined (Figures 4g-l). The pseudostomal 
aperture was invaginated about /4 of the test height and 
its structure included a well-developed buccal tube 
(Figures 4b, f). Of all measurements made on tests of 


'After submission of this paper by K. Nicholls to The Canadian Field-Naturalist in February, 2005, one of the referees selected by the editor 
to review it (R. Meisterfeld) informed Nicholls that A. formosa had been previously found by him in Germany and by J. To6r6k in Hungary. 
Although neither discovery had been submitted for publication, a poster presentation by Térék had been made in Italy at the 4° European 
Congress of Protistology under the unofficial name, “Arcella siemensmai’. The inclusion of Meisterfeld and Torok as co-authorities of A. for- 
mosa in this paper was agreed by all as an acceptable way to acknowledge the original and preemptory submission by Nicholls as well as the 
independent discoveries of this taxon by Meisterfeld and Torék. As a consequence, the names “A. siemensmai” and “A. robusta” previously 
used by Tor6k & Meisterfeld and F. Siemensma, respectively, to describe this taxon are rendered invalid. 


2005 


NICHOLLS: Two NEW SPECIES OF TESTATE RHIZOPODS 


407 


FIGURE 3. Pseudostomal apertures in nine different specimens of Cyclopyxis acmodonta illustrating the wide range in shape. 


The scale bar in Figure 3a applies to all Figures 3a-i. 


this species, the greatest variation was found in the 
aperture diameter (coefficient of variation = 24.5%; 
Table 1). 

The test of A. formosa appears to be of two major 
structural components: an underlying meshwork of 
irregularly sized and shaped meshes (Figure 5a), and 
an overlying sheet of elliptical-to-circular disc-like are- 
oles averaging about 3 x 4 um. Small, but well-defined 
pores are located at the junctions of adjacent areoles 
so that each areole is surrounded by usually 4-5 pores 
(Figure 5b). This two-layered structure was revealed 
in the broken test of a specimen that apparently had a 
large piece of the outer areolar layer stripped off (Fig- 
ure 5a; but note the small patch containing the surface 
areolar layer in the lower right of Figure 5a). 


Discussion 

The genus Cyclopyxis contains a few large species 
with lobed pseudostomal apertures; consequently, the 
test structure of each had to be reviewed before any 
conclusions about the autonomy of C. acmodonta was 
established. Cyclopyxis impressa (Daday) Deflandre 
(= Difflugia lobostoma var. impressa Daday = Cen- 
tropyxis impressa (Daday) Da Cunha) is apparently 


restricted to a few locations in the Southern Hemi- 
sphere (Velho et al. 1996). All reports show a much 
larger test (300-561 tm in diameter) than that found 
for C. acmodonta; other differences include a very 
regularly shaped and symmetrical pseudostomal aper- 
ture with 5-8 lobes, a much greater diameter-to-height 
ratio, and a much greater degree of invagination of the 
pseudostome (to about 50% of the test height). Cyclopy- 
xis trilobata var. maxima Chardez, 1971, test size 
(D = 225-235; H = 110-140 um) is close to C. acmod- 
onta. Although Chardez (1971) did not illustrate his new 
taxon (to which I attribute subspecies status implied 
by his term “var.”’), he stated that C. triblobata var. 
maxima differed from C. trilobata Bartos,1963 only 
in its much greater size. Like its nominotypical sub- 
species, Cyclopyxis trilobata var. maxima has a small, 
well-defined, three-lobed pseudostome which clearly 
distinguishes it from C. acmodonta. 

Bartos (1963) described Cyclopyxis crucistoma 
which is significantly smaller than C. acmodonta with 
a test diameter of 122-124 um and a height of 50 um. 
It also has a well-defined pseudostome in the shape 
of a cross. Cyclopyxis grospietschi (Sch6nborn 1962) 
also has a pseudostome in the shape of a cross, but its 


408 


test diameter and height are only 125 and 80 um, 
respectively (significantly smaller than C. acmodonta). 
Decloitre (1954) described Cyclopyxis lobostoma with 
a 7-lobed pseudostome and a test diameter and height 
of 430 and 280 um, respectively according to Bartos 
(1963), or 400 and 300 um, according to Decloitre 
(1977). In ventral view, this species resembles C. im- 
pressa, but its pseudostome is apparently not invaginat- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


ed. This fact is grounds for questioning its placement 
in the genus Cyclopyxis, so it needs to be rediscovered 
and evaluated relative to other genera in the Trigonopy- 
xidae (possibly a Geopyxella species?). 

As regards overall test shape and degree of invagi- 
nation of the pseudostome, C. acmodonta’s closest “‘tel- 
ative” would appear to be C. stellata (Wailes 1927) Defi. 
1929. Important differences between the two species 


FIGURE 4. Arcella formosa tests. a. Drawing of the ventral surface (translucency of the test allows the ridges separating the 
depressions on the dorsal surface to be revealed; see also Figure 4j; scale bar applies to all Figures 4a-l). b. Drawing of 
the lateral view; the pseudostomal aperture (ap) is on the ventral surface; D = test diameter; H = test height. c. Image 
of the ventral view of a test. d. Image of the lateral view of the same specimen illustrated in Figure 4c. e. Image of 
the ventral view of a different specimen. f. Image of the lateral view of the same specimen illustrated in Figure 4e; 
arrows indicate the location of the buccal tube. g-i. Images of the lateral view of a different specimen at three differ- 
ent levels of microscopic focus. j-l. Images of the ventral side (j), sub-dorsal (k), and dorsal (1) surfaces of the same 
specimen illustrated in Figures 4 g-i. 


2005 


are as follows: (1) C. stellata has a pseudostomal aper- 
ture with 3-5 well-defined lobes while C. acmodonta’s 
aperture is irregularly shaped. (2) The distinctive sharp- 
pointed granules ornamenting the internal margin of the 
pseudostome in C. acmodonta are apparently not pres- 
ent in C. stellata. (3) With test diameters and heights 
of 335-400 and 252-290 um, respectively, C. stellata 
is significantly larger than C. acmodonta; the smallest 
C. stellata tests are larger than the biggest C. acmod- 
onta tests. (4) Wailes (1927) described the test of C. 
stellata as being “composed of irregularly shaped sili- 
ceous plates, without protuberances”. In C. acmodonta, 
although there were some flat plate-like particles seen, 
the test elements are more aptly described as “irregu- 
larly shaped polygonal particles”. 

In a review of the variability and taxonomy of sev- 
eral smaller species of Cyclopyxis, Foissner and Korga- 
nova (1995) found a wide range of sizes among some 
species. They concluded that size criteria may be of 
limited value in distinguishing among certain species 
unless the differences are very distinct, and/or the size 
difference is accompanied by at least one other reli- 
able morphologic character. It is not known how such 
criteria might apply to the larger Cyclopyxis species 
with lobed pseudostomal apertures, because measure- 
ments of large numbers of specimens and the appro- 
priate follow-up statistical analyses have not been 
published for most. With the literature data available, 
however, C. acmodonta’s test size and other morpho- 
metric features (pseudostome shape and degree of 
invagination) clearly set this species apart from other 
similar Cyclopyxis species. 

Unfortunately the nomenclature of the three Cen- 
tropyxis species (C. stellata Wailes, C. arcelloides 
Pénard and C. impressa (Daday) Da Cunha) transferred 
to Cyclopyxis by Deflandre (1929) is somewhat con- 
fused. Undoubtedly this confusion stems from Deflan- 
dre’s own treatment of Cyclopyxis in later years. Since 
the erection of Cyclopyxis in 1929, virtually all stu- 
dents of the Arcellinida have considered Cyclopyxis a 
separate and well defined genus (Decloitre (1977). 
Deflandre’s post-1929 treatment of Cyclopyxis, how- 
ever, remains ambiguous. Deflandre (1953) acknowl- 
edged that other authors had treated Cyclopyxis and 
Centropyxis as separate and distinct genera, but by this 
date (1953) Deflandre himself apparently still had not 
accepted this. He did not list Cyclopyxis anywhere in 
his classification of the Testacealobosa, except noting 
its existence in a footnote to the genus Centropyxis, 
within which he continued to submerge it. Several 
years later, in his comprehensive summary and key 
of freshwater testate rhizopods, Deflandre (1959) did 
not list any Cyclopyxis species but named two Cyclopy- 
xis species under Centropyxis (Centropyxis stellata 
Wailes and Centropyxis arcelloides Penard). 

Other sources of confusion include (1) Harnisch 
(1958), who considered Cyclopyxis a subgenus of 
Centropyxis, and, like Deflandre (1959) listed “Cen- 
tropyxis stellata Wailes”, and (2) Chardez (1967) who 


NICHOLLS: Two NEw SPECIES OF TESTATE RHIZOPODS 


409 


FiGuRE 5. Microscopic structure of the test wall of Arcella 
formosa. a. Underlying meshwork of test wall showing 
variation in shape and size of meshes. Small black 
arrow points to the interior rim of the pseudostomal 
aperture with its palisade layer of meshes. Larger 
white arrow points to a small patch of overlying sub- 
circular areoles. b. Piece of broken test wall showing 
the arrangement of circular to elliptical areoles with 
interspersed pores. Scale bar applies to both Figures 
5a and b. 


listed ““Cyclopyxis stellata Wailes” despite the fact that 
Wailes had described it as a Centropyxis species and 
that it had not previously been formally assigned to 
Cyclopyxis as anew combination. Chardez (1967) listed 
Centropyxis and Cyclopyxis as distinct genera in the 
family Centropyxidae, but listed 12 other genera, 11 
of which had not been included in Deflandre’s (1953) 
original concept of the family. Chardez (1967, and later 
papers) did not formally revise the description of the 
Centropyxidae to include the broadened range of form 
implied by inclusion of the 11 additional genera. 
Although Jung (1942) first introduced the family 
name Centropyxidae, with the result that Jung (1942) 
is sometimes listed as the authority for this family (e.g., 
Bovee 1985; Meisterfeld 2002), the correct authority 
is Deflandre (1953) who first provided a formal cir- 
cumscription and included four genera. More recently, 
however, the family Centropyxidae has implicitly be- 
come better defined with the recognition that several 
genera originally on Chardez’s (1967) list of Centro- 
pyxidae could be more naturally accommodated within 
the family Trigonopyxidae Loeblich and Tappan, 1964. 
In the future, in order to correct some of the prob- 
lems outlined above, students of the taxonomy of these 
arcellinid families should list the authority for the Cen- 


410 


tropyxidae as follows: “Centropyxidae Deflandre, 1953 
sensu Meisterfeld 2002”, in order to reflect its con- 
temporary generic composition. As well, species trans- 
ferred from Centropyxis to Cyclopyxis by Deflandre 
(1929) should be rendered in accordance with Article 
51 of the Interntional Code of Zoological Nomencla- 
ture (ICZN) as a combination attributed to Deflandre; 
e.g., Cyclopyxis arcelloides (Penard) Deflandre, as list- 
ed in Jung (1942) and Laminger (1972), for example, 
and not as Cyclopyxis arcelloides Penard, as listed in 
Chardez (1967), Bonnet (1977), and Cotiteaux and 
Chardez (1981). Species of Cyclopyxis described after 
Deflandre (1929) should not pose any nomenclatural or 
authorship difficulties because no new combinations 
are required (e.g., Cyclopyxis crucistoma Bartos, 1963). 

There are virtually no Arcella species that can be 
confused with A. formosa. Firstly, its large size is high- 
ly unusual among representatives of the genus. Only A. 
artocrea Leidy emend. Deflandre, A. rota Daday, A. 
megastoma Penard, A. leidyana Deflandre and A are- 
naria var. grandis Bunescu & Matis are of comparable 
size. All of the above-listed species have markedly dif- 
ferent test shapes including much greater or lesser test 
diameter-to-height ratios, much smaller pseudostomal 
apertures of different structure (e.g., greater degree of 
invagination on the ventral surface, presence of large 
pores surrounding the aperture rim). 

Many Arcella species have dorsal surfaces ornament- 
ed with shallow depressions separated in some cases by 
thickened ridges on the test wall. Some of these include 
A. crenata Playfair, A. bathystoma Deflandre, A. arto- 
crea Leidy ssp. pseudocatinus Deflandre and A. gib- 
bosa Penard, among several others. Again, these all 
have major size and structural differences that clearly 
set them apart from A. formosa. The degree of form 
variation in A. formosa was objectively quantified in 
14 randomly encountered specimens and subjectively 
evaluated in several more specimens so that there can 
be little doubt that its morphology is distinct and sep- 
arate from any previously described species. 

Species that are very small in size likely have a great 
potential to be overlooked in investigations of testate 
rhizopods. When new species of small-sized taxa are 
discovered, conclusions about geographic distribution 
may not generally be possible because of the potential 
for such species to exist in many habitats over large 
geographic areas, but to have escaped previous detection 
because of their small size. Because of their large sizes, 
both A formosa and C. acmodonta, on the other hand, 
were readily detected in the Ontario samples. Recog- 
nizing that many species of Arcella and Cyclopyxis 
(earlier as Centropyxis) were known to science nearly 
a century ago, if A. formosa and C. acmodonta were 
widespread in other habitats in other parts of the world, 
they should have been detected by others long ago 
(notwithstanding the recent discoveries of C. acmo- 
donta in Hungary and Germany: (R. Meisterfeld, per- 
sonal communication). The logical conclusion is that 
Cranberry Marsh and the Beaver River coniferous for- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


est bogs afford these species certain environmental tol- 
erances or requirements not widely available elsewhere. 
This lends support to the widely held view among local 
naturalists and biologists familiar with the botanical and 
avian attributes of these habitats that these are areas 
of unique biological status deserving of special pro- 
tection and further scientific investigation. 


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Scott, L. D., J. G. Winter, M. N. Futter, and R. E. Girard. 
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Chardez, D., L. Beyens, and P. De Bock. 1987. Arcella ovali- 
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Cotiteaux, M-M., and D. Chardez. 1981. Thécamoebiens 
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Decloitre, L. 1979. Mises a jour au 31.12.1978 des mises a 
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Decloitre, L. 1982. Compléments aux publications précé- 
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Received 11 February 2005 
Accepted 15 August 2005 


A Range Extension for the Rock Vole, Microtus chrotorrhinus, 


in Labrador 


SARAH W. LANSING 


Division of Vertebrate Natural History, Michigan State University Museum, Michigan State University, East Lansing, Michigan 


48824 USA; e-mail: Sarah.Lansing @asu.edu 


Lansing, Sarah W. 2005. A range extension for the Rock Vole, Microtus chrotorrhinus, in Labrador. Canadian Field-Naturalist 


119(3): 412-416. 


Rock Voles (Microtus chrotorrhinus) were previously documented only from the southern coast of Labrador and the Strait 
of Belle Isle south of Hamilton Inlet. During two field seasons in 2000 and 2001, both in July, six Rock Vole specimens were 
collected on Southern Island, off the north-central coast of Labrador, extending the range of the subspecies Microtus chrotor- 
rhinus ravus approximately 250 km to the central Labrador coast north of Hamilton Inlet. Rock Voles were trapped primarily 
in stunted Black Spruce (Picea mariana) forest and areas associated with rocks, thick brush, and water. 


Key Words: Rock Vole, Microtus chrotorrhinus, Labrador, distribution. 


Rock Voles (Microtus chrotorrhinus) occur from the 
Ungava Peninsula of Labrador to the southeastern 
United States, and west into Minnesota, but their distri- 
bution in Labrador is not well documented. Two sub- 
species of Microtus chrotorrhinus are currently recog- 
nized in Canada: M. chrotorrhinus chrotorrhinus, 
distributed widely across eastern Canada and into the 
midwestern portions of the United States (Anderson 
1946; Banfield 1974; Kirkland and Jannett 1982; 
Komarek 1932) and M. chrotorrhinus ravus, recorded 
only from the Ungava Peninsula of Labrador and ap- 
parently restricted to the coastal regions (Anderson 
1946; Banfield 1974; Kirkland and Jannett 1982; 
Komarek 1932). In this study, Rock Voles were trapped 
in the Adlavik Islands, Labrador, and are most likely 
identified to M. chrotorrhinus ravus, which is known 
from the type locality of Black Bay, Strait of Belle Isle, 
and from L’Anse-au-Loup, Red Bay, Mary’s Harbour, 
and Hare Harbour, Labrador (Anderson 1946; Harper 
1961; Peterson 1962; Whitaker and Martin 1977). Ban- 
field (1974) documented M. chrotorrhinus ravus on 
Anticosti Island in the Gulf of St. Lawrence, but 
Cameron (1958) did not collect this subspecies on the 
island. To my knowledge, the only confirmed island 
population of Rock Voles has been reported from Cape 
Breton Island, Nova Scotia (Roscoe and Majka 1976). 
Thus, not only does this study extend the range of Rock 
Voles farther north in Labrador but also documents a 
second population of Rock Voles on an island. 

Throughout their range, Rock Voles are generally 
associated with habitats that feature rocks or talus 
slopes (Kirkland 1977; Kirkland and Knipe 1979; 
Roscoe and Majka 1976; Timm et al. 1977). Rock Voles 
usually occur in moist mossy areas near streams and 
ponds, thick brush, and open-canopy forests (Buech et 
al. 1977; Kirkland and Jannett 1982; Kirkland and 
Knipe 1979), although Nagorsen and Peterson (1981) 
collected Rock Voles on ridges of dry upland conifer- 


ous forest near lake margins. Groundcover and berry 
plants are an important component of Rock Vole micro- 
habitats. Rock voles are often trapped in association 
with important food sources, including Labrador Tea 
(Rhododendron groenlandicum), Bunchberry (Cornus 
canadensis), Bakeapple or Cloudberry (Rubus chamae- 
morus), Alpine Bilberry (Vaccinium uliginosum), and 
Partridgeberry (Vaccinium vitis-idaea) (Kirkland and 
Jannett 1982; Timm et al. 1977; Whitaker and Martin 
1977). Roots, seeds, grass stems and leaves, and larvae 
of insects have also been recorded in the stomach con- 
tents of Rock Voles (Whitaker and Martin 1977). 


Study Area 

During two field seasons in Labrador, surveys of 
small mammals were conducted in the vicinity of Ad- 
lavik Harbour (55°O1'N, 58°49"W; Datum: NAD 27; 
see also Canada, Surveys and Mapping Branch 1964a,b) 
on Southern Island in the Adlavik Islands (Figure 1). 
The Adlavik Islands consist of 11 primary landmasses 
and are off the north-central coast of Labrador, approx- 
imately 25 km south of the small coastal community 
of Makkovik (Figure 1). From the most southern point 
on Southern Island to the nearest point on the mainland 
is 3.75 km. Trapping sites are approximately 10 km 
from the mainland. During the winter, Southern Island 
is connected to the mainland by ice, which would allow 
small mammals to traverse from the mainland to the 
islands. 

The Adlavik Islands are characterized by stands of 
stunted black spruce (Picea mariana) forest that shel- 
ter dwarf thickets and brush, carpets of moss and lichen, 
and rocky tundra on the upper slopes and summits of 
the hills; habitat that is consistent with previous des- 
criptions of vegetational communities along the central 
Labrador coast (Fitzhugh 1972; Tanner 1947). The 
Ecological Stratification Working Group (1996) places 
the Adlavik Islands in the taiga shield ecozone, specif- 


412 


2005 


ically the coastal barrens ecoregion, which is charac- 
terized by long cold winters and short wet summers. 
Mean temperatures range from —13.5°C in the winter 
to 7°C in the summer and annual precipitation can 
range from 600-1000 mm (Ecological Stratification 
Working Group 1996). The coastal barrens are part of 
the Atlantic low subarctic ecoclimatic region and are 
dominated by stands of black spruce and understories 
of Dwarf Birch (Betula sp.), Labrador Tea, Lichen 
(Cladina sp.), and moss (Ecoregions Working Group 
1989). 


Methods and Results 

Small mammals were trapped from 11 July to 27 
July 2000 and from 4 July to 22 July 2001. Trapping 
sites were located in representative vegetational com- 
munities on the hillside above Adlavik Harbour. During 
the 2000 field season, five traplines were set 0.08 km 
SW of Adlavik Harbour (55°01.426'N, 58°49.050'W, 
Datum: NAD 27) at an elevation between 15 and 20 m 
above sea level. Four traplines with 10 traps each ran 
vertically up the hill; two traplines with five and four 
traps, respectively, were initially located on open, moist, 


59° 


BR 
SF 


oT ie 


55” 


LANSING: ROCK VOLE IN LABRADOR 


413 


grass-covered rocks of the lowland coastline, approx- 
imately 3 m above sea level. After two days, these latter 
two traplines were condensed into one trapline now 
with 10 traps and positioned horizontally across the hill 
near the other lines, as specimens were not trapped 
along the coastline. Each of the five traplines encom- 
passed multiple microhabitats such that 19 traps were 
set in dense microhabitat (thick stunted spruce forest 
microhabitat with numerous water sources), 27 traps in 
less dense microhabitat (typically more open bush 
microhabitat with thick brush, mosses and lichens, and 
rocks), and four traps in open microhabitat (microhab- 
itat with dry, gravelly soil beneath boulders with little or 
no brush). Traps were not set along the summit of the 
hill, which was only sparsely vegetated. 

During the 2001 field season, five traplines (0.15 km 
SW of Adlavik Harbour, 55°01.419'N, 58°48.976'W) 
were set adjacent to the trapping area of the 2000 
field season at an elevation between 15 and 40 m above 
sea level. Four traplines were positioned horizontally 
across the hill and 1 line ran vertically up the hill. After 
five days of trapping, one trapline was moved to the 
rocky knoll next to camp (0.56 km NE of Adlavik 


Adlavik 


2 


ip aels 

oid) i 

- e 
Os 


50° 
eg, Oe 
“55° =54> 


-57° -56° 


-53° 


-52° 


FIGURE 1. Map showing the previously documented range (gray shading) of Rock Voles (Microtus chrotorrhinus) in Labrador 
and the current trapping results in the vicinity of Adlavik Harbour, Southern Island. 


414 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |. Total length (TL), tail vertebra length (TV), hindfoot length (HF), ear length (EAR), and weight (WT) of male and 


female Microtus chrotorrhinus from Southern Island. 


TL TV 

Adult Female 

FMNH 168637 149 44 
FMNH 168638 149 50 
FMNH 176626 127 41 
Mean 141.67 45 
Sub-adult Female 

FMNH 168640 132 35 
Juvenile Male 

FMNH 168639 119 35 
FMNH 168641 118 40 


Harbour; 55°01.757'N, 58°49.217'W; 10 m) and 2 of 
the remaining lines were shifted to the same spruce 
and open bush habitat sites that were trapped in 2000 
and were oriented vertically up the hill. From 4 July 
to 8 July 2001, traps were set in dense spruce micro- 
habitat (n=11), in less dense bush microhabitat (n=33), 
and in open habitat of boulders and gravel (n=6). From 
11 July to 22 July 2001, traps were located in dense 
spruce microhabitat (n=22), in less dense bush micro- 
habitat (n=14), and in open microhabitat (n=14). All 
10 traps on the rocky knoll above camp were on a talus 
slope in open microhabitat with boulders and gravel. 
Each trapline usually consisted of 10 Museum Spe- 
cial traps spaced at 5-m intervals; traplines were approx- 
imately 10 m apart. Traps were placed in different 
microterrains, including beneath rock overhangs, beside 
logs, along runways in the vegetation, and under small 
bushes and brush. If traplines were vertical to the slope 
of the hill, traps were also vertically oriented and vice 
versa for horizontally positioned traplines. Traps were 
baited with a mixture of peanut butter and rolled oats 
and checked twice per day, once in the early morning 
and once in the early evening. Specimens were prepared 
as skins and skeletons or as fluid specimens in the field. 
Skeletons and skulls were cleaned by a dermestid beetle 
colony at the Field Museum of Natural History. Cleaned 
skulls were measured, following the methods in Martin, 
Pine, and DeBlase (2001), using Mitutoyo digital cali- 
pers to the nearest 0.01 mm. Specimens are deposited 
at the Field Museum of Natural History in Chicago, 
Illinois, USA (museum numbers: 2000 — FMNH 
168599-168649, 2001 — FMNH 176624-176629). 
During the 2000 field season, 51 small mammals, 
representing six species were trapped: Pygmy Shrew 
(Sorex hoyi, n=10), Deer Mouse (Peromyscus manic- 
ulatus, n=7), Southern Red-backed Vole (Clethriono- 
mys gapperi, n=21), Rock Vole (n=5), Meadow Vole 
(Microtus pennsylvanicus, n=1), and Heather Vole 
(Phenacomys intermedius, n=7). During trapping in 
2000, four Rock Voles were found in dense habitats of 
stunted spruce forest, thick brush, mosses, lichens, and 
berry plants with intermittent water sources and one 
was in relatively thick brush near trickling water. Three 
Rock Voles were trapped beneath rock overhangs and 


HF EAR WT (g) 
20 18 33 
21 16 39 
21 15 19 
20.67 16.33 30.33 
21 fe) 20 
20 13 15 
22 15 Pi) 


two beneath bushes in more open habitat. The Rock 
Voles (n=5) obtained include adult females (n=2), sub- 
adult females (n=1), and juvenile males (n=2). Two 
female Rock Voles displayed large mammae, perforated 
vaginas, and no embryos (FMNH 168638 and 168640). 
One female had small mammae, an imperforated vagi- 
na, and no embryos (FMNH 168637). Two males had 
scrotal testes (FMNH 168639: testes size 7 mm long by 
5 mm wide; FMNH 168641: testes size 11 mm long by 
6 mm wide). 

During the 2001 field season, only 6 individuals were 
trapped: Deer Mouse (n=5) and Rock Vole (n=1). The 
single Rock Vole was trapped in dense habitat of stunt- 
ed spruce forest beneath a rock overhang and a stunted 
spruce. This individual was an adult female with small 
mammae, a perforated vagina, and no embryos. 

Body measurements of Rock Voles from different 
age classes are presented in Table 1. Several other stud- 
ies have reported similar body measurements for this 
species (Banfield 1974; Kirkland 1977; Kirkland and 
Jannett 1982; Komarek 1932; Timm 1974; Timm et al. 
1977). Cranial measurements of adult female Rock 
Vole specimens are presented in Table 2. Skulls were 
measured only if all sutures were fused, indicating 
adult status. Komareck (1932) reports average cranial 
measurements for seven specimens of Rock Voles of 
unknown sex and Timm (1974) documents select cra- 
nial measurements for a male and female Rock Vole, 
which are similar to the sampled Adlavik specimens. 
Although body and cranial measurements for the adult 
female Rock Vole trapped in 2001 are small in com- 
parison to the two adult females from 2000, the spec- 
imen exhibits equivalent fusion of cranial sutures, 
degree of toothwear, and closure of dentine triangles on 
molars. Since the small mammal population appears 
to have declined on Southern Island in 2001, changes 
in resource availability could have contributed to the 
small size of the adult female Rock Vole trapped dur- 
ing that field season. 


Discussion 

In this study, Rock Voles were trapped in rocky wet 
habitats associated with dense cover or under thick 
bushes in more open areas. As noted by most authors 


2005 LANSING: ROCK VOLE IN LABRADOR 415 


TABLE 2. Cranial measurements of three adult female Microtus chrotorrhinus from Southern Island. 


Skull Measurements FMNH 168637 FMNH 168638 FMNH 176626 Range 

Greatest length of skull 24.57 24.83 pH IG iP) 23.72 - 24.83 
Condylobasal length 24.87 253 23.67 23.67 - 25.15 
Least interorbital breadth 3.74 4.01 3.92 3.74- 4.01 
Nasal length 741 ie 6.86 6.86 - 7.41 
Palatal length 1255 12.82 12.44 12.44 - 12.82 
Mastoid breadth 10.92 10.88 10.21 10.21 - 10.92 
Zygomatic width 13.67 14.44 12.92 12.92 - 14.44 
Basilar length 22.02 22.24 21.01 21.01 - 22.24 
Basal length jhe 23.64 22.54 22.54 - 23.64 
Maxillary tooth row 3:67 6.00 5.64 5.64 - 6.00 
Mandible length L553 14.14 12.74 12.74 - 14.14 


(for an exception, see Nagorsen and Peterson 1981), 
across their range Rock Voles prefer rocky densely cov- 
ered environments with abundant water sources (Kirk- 
land and Jannett, 1982; Kirkland and Knipe, 1979; 
Timm et al., 1977). The Ecological Stratification Work- 
ing Group (1996) notes that steep talus slopes, usually 
a component of Rock Vole habitat, are a common 
feature of the coastal barrens ecoregion. 

The winter prior to the 2001 survey was very harsh, 
exceptionally cold and snowy, which could explain 
the paucity of small mammals trapped that season. 
Voles are known to cycle through high and low popu- 
lation densities, although the causes of cyclic popula- 
tion fluctuations are still debated. Pruitt (1972) docu- 
mented synchronous fluctuations in small mammal 
populations on the island of Newfoundland and main- 
land Labrador, which are separated by the Strait of 
Belle Isle. Pruitt (1972) suggests that changes in eco- 
system productivity could be the primary factor con- 
trolling population fluctuations. 

Prior to this study Rock Voles were documented 
only from south of Hamilton Inlet in Labrador and from 
one coastal island. The surveys on Southern Island 
extend the range of Rock Voles approximately 250 km 
north of Hamilton Inlet and provide the second record 
of Rock Voles on a coastal island. Southern Island is 
located in close proximity to mainland Labrador and 
ice connects both landmasses during the winter such 
that small mammals could become established on 
islands. Since many of the Adlavik Islands are not far 
apart and close to mainland and share similar habitat 
features, Rock Voles could be expected to occur on 
other islands. Future sampling in Labrador could 
expand the known distribution of Rock Voles farther 
north in Labrador and provide further data on diet, 
habitat preferences, and measurements. 


Acknowledgments 

The Inland Fish and Wildlife Division of the Depart- 
ment of Tourism, Culture, and Recreation of Newfound- 
land and Labrador granted permission for this study. I 


am most grateful to Robert Otto, Labrador Senior Wild- 
life Biologist of the Inland Fish and Wildlife Division, 
for permission to trap small mammals and for encour- 
aging research by an undergraduate student. Trapping 
equipment and supplies were provided by the Field 
Museum of Natural History, Division of Mammals. I 
thank Bill Stanley for all his help in field preparations 
and specimen identification. These surveys simulta- 
neously occurred with archaeological fieldwork, direct- 
ed by Stephen Loring and Leah Rosenmeier, at Adlavik 
Harbour. Many thanks to Leah Rosenmeier for con- 
structing a map for this paper. I thank Stephen Loring, 
Leah Rosenmeier, Larry Heaney, and the Adlavik Island 
field crew for their support and many useful sugges- 
tions. [ am very grateful to Bill Stanley, Barbara Lun- 
drigan, Stephen Loring, and Russ Van Horn for their 
comments on previous drafts of this paper. I also wish 
to thank David Nagorsen and two anonymous review- 
ers for very helpful revisions of this manuscript. 


Literature Cited 

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mals. National Museum of Canada Bulletin 102 (Biological 
Sciences Series 31). 

Banfield, A. W. F. 1974. The mammals of Canada. University 
of Toronto Press, Toronto. 

Buech, R. R., R. M. Timm, and K. Siderits. 1977. A second 
population of rock voles, Microtus chrotorrhinus, in Min- 
nesota with comments on habitat. Canadian Field-Naturalist 
91: 413-414. 

Cameron, A. W. 1958. Mammals of the islands in the Gulf of 
St. Lawrence. National Museum of Canada Bulletin 154: 
1-165. 

Canada, Surveys and Mapping Branch. 1964a. Rigolet, 
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Canada, Surveys and Mapping Branch. 1964b. Makkovikt, 
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Ecological Stratification Working Group. 1996. A national 
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Ecoregions Stratification Working Group. 1989. Ecocli- 
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Fitzhugh, W. W. 1972. Environmental archaeology and cultur- 
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Harper, F. 1961. Land and freshwater mammals of the Unga- 
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Kirkland, G. L., Jr. 1977. The rock vole, Microtus chrotorrhi- 
nus (Miller) (Mammalia: Rodentia) in West Virginia. Annals 
of the Carnegie Museum 46: 45-53. 

Kirkland, G. L., Jr., and F. J. Jannett, Jr. 1982. Microtus 
chrotorrhinus. Mammalian Species 180: 1-5. 

Kirkland, G. L., Jr., and C. F. Knipe. 1979. The rock vole 
(Microtus chrotorrhinus) as a transition zone species. Cana- 
dian Field-Naturalist 93: 319-321. 

Komarek, E. V. 1932. Distribution of Microtus chrotorrhinus, 
with description of a new subspecies. Journal of Mammalo- 
gy 13: 155-158. 

Martin, R. E., R. H. Pine, and A. F. DeBlase. 2001. A manual 
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Nagorsen, D. W., and R. L. Peterson. 1981. Distribution, 
abundance and species diversity of small mammals in 
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Roscoe, B., and B. Majka. 1976. First records of the rock 
vole (Microtus chrotorrhinus) and the Gaspe shrew (Sorex 
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history of rock voles (Microtus chrotorrhinus) in Minneso- 
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Received 29 December 2003 
Accepted 20 July 2005 


New Records of Vascular Plants in the Yukon Territory VII 


WILLIAM J. Copy!, BRUCE A. BENNETT’, and PHIL CASWELL? 


| Biodiversity, National Program on Environmental Health, Agriculture and Agri-Food Canada, Wm. Saunders Building (49), 
Central Experimental Farm, Ottawa, Ontario K1 A 0C6 Canada 

Department of Environment, Government of the Yukon, Box 2703, Whitehorse, Yukon Y1A 2C6 Canada; e-mail: Bruce. 
Bennett @ gov.yk.ca 

deceased 


i) 


William J. Cody, Bruce A. Bennett, and Phil Caswell. 2005. New records of vascular plants in the Yukon Territory VII. Cana- 
dian Field-Naturalist 119(3): 417-436. 


Based on field reconnaissance mainly in 2003 in Kluane National Park but also many other areas throughout the Yukon, 
information is provided on geographically significant plant occurrences. Thirteen native taxa, Agrostis humilis, Blysmopsis 
rufa, Carex bebbii, C. lenticularis var. dolia, C. nigricans, C. sartwellii, C. siccata, Diphasiastrum sitchense, Najas flexilis, 
Oxytropis arctica var. arctica, Oxytropis arctica var. murrayi, Saxifraga nelsoniana ssp. carlottae and Swertia perennis and 
two introduced taxa, Bromus japonicus and Leymus mollis ssp. mollis s.\. are reported new to the flora of the Yukon Territory. 
Significant range extensions for 109 native and eight introduced taxa are included and three species, Clematis occidentalis, 


Polygonum persicaria, and Verbena hastata are deleted from the flora. 


Key Words: Vascular plants, Yukon Territory, flora, new records, range extensions, phytogeography. 


Since the writing of New Records of Vascular Plants 
in the Yukon Territory VI (Cody et al. 2004), a consid- 
erable number of plant specimens have been submitted 
to Cody for identification and confirmation. The major 
submissions include the following locations and col- 
lectors: (1) Kluane National Park and vicinity by Phil 
Caswell and Bruce A. Bennett for the National Park 
Service as part of ongoing botanical inventories; (2) 
various locations by Reidar Elven and Heidi Solstad 
visiting from the University of Oslo, Norway; (3) 
Bruce A. Bennett collected plant specimens from 
throughout the territory; and (4) Greg Brunner from 
the Dawson City area. 

This paper serves to further update the Flora of the 
Yukon Territory (Cody 1996) and Flora of the Yukon 
Territory, Second Edition (Cody 2000) along with other 
records recently published (Cody et al. 1998, 2000, 
2001, 2002, 2003, 2004). The floristic information 
presented earlier and updated here provides the basis 
for biological research and ongoing work relating to 
wildlife management, forestry, sustainable resource 
management and agriculture. With additions of thir- 
teen native and two introduced species reported here 
the flora now includes 1195 species. 

The taxa addressed in the body of this paper appear 
in a synoptic list by Yukon status in alphabetical order. 
The taxa are then discussed in taxonomic order, as pre- 
sented in the Flora of the Yukon Territory with citation 
of specimens and other pertinent information. Com- 
mon names follow Cody (1996), Douglas et al. (1998- 
2001) and Kartesz and Meacham (1999). 


Synoptic List by Yukon Status 

Native Taxa New to the Yukon Territory (13): 
Agrostis humilis 

Blysmopsis rufa (Scirpus rufus) 


Carex bebbii 

Carex lenticularis var. dolia 

Carex nigricans 

Carex sartwellii 

Carex siccata 

Diphasiastrum sitchense (Lycopodium sitchense) 
Najas flexilis 

Oxytropis arctica var. arctica 
Oxytropis arctica var. murrayi 
Saxifraga nelsoniana ssp. carlottae 
Swertia perennis 


Introduced Taxa New to the Yukon Territory (2): 
Bromus japonicus 
Leymus mollis ssp. mollis s.1. 


Range Extensions of Native Taxa within the 
Yukon Territory (109): 
Angelica lucida 

Antennaria pulcherrima 
Aphragmus eschscholtzianus 
Apocynum androsaemifolium 
Arabidopsis salsuginea 

Arabis boivinii 

Arnica diversifolia 

Arnica latifolia 

Artemisia tilesii 

Astragalus alpinus 

Athyrium filix-femina var. cyclosorum 
Campanula rotundifolia 

Carex brunnescens 

Carex buxbaumii 

Carex canescens 

Carex eburnea 

Carex lasiocarpa 

Carex laxa 

Carex lenticularis var. lipocarpa 
Carex livida 

Carex microglochin 

Carex microptera 


417 


418 THE CANADIAN FIELD-NATURALIST 


Carex nardina 

Carex parryana 

Carex phaeocephala 

Castilleja miniata 

Castilleja parviflora 

Castilleja unalaschcensis 

Cirsium foliosum 

Coeloglossum viride ssp. bracteatum 
Comandra umbellata ssp. pallida 
Cryptogramma crispa Vat. sitchensis 
Douglasia ochotensis 

Draba albertina 

Draba cinerea 

Draba crassifolia 

Draba lonchocarpa vat. vestita 
Draba nivalis 

Draba oligosperma 

Draba scotteri 

Draba stenoloba 

Draba stenopetala 

Drosera anglica 

Dryas integrifolia ssp. crenulata (Dryas crenulata) 
Dryas octopetala ssp. hookeriana (Dryas hookeriana) 
Epilobium hornemannii ssp. hornemannii 
Erigeron grandiflorus ssp. arcticus 
Erigeron humilis 

Erigeron peregrinus ssp. peregrinus 
Erigeron pumilus 

Erigeron uniflorus ssp. eriocephalus 
Erigeron yukonensis 

Eriophorum gracile 

Festuca brachyphylla 

Festuca brevissima 

Festuca minutiflora 

Galium triflorum 

Geranium erianthum 

Harrimanella stellariana 
Hieracium gracile 

Iris setosa ssp. interior 

Juncus bufonius 

Juncus filiformis 

Leptarrhena pyrolifolia 

Luetkea pectinata 

Lupinus nootkatensis 

Luzula piperi 

Luzula spicata 

Menyanthes trifoliata 

Minuartia dawsonensis 
Myriophyllum verticillatum 

Nuphar variegatum 

Orobanche fasciculata 

Oxytropis campestris ssp. jordalii 
Oxytropis nigrescens ssp. nigrescens 
Papaver radicatum ssp. kluanensis 
Parnassia fimbriata 

Phippsia algida 

Phyllodoce X intermedia 

Pinus contorta ssp. latifolia 
Plantago eriopoda 

Polemonium pulcherrimum 
Polygonum achoreum 

Polygonum lapathifolium 
Polystichum lonchitis 

Potamogeton praelongus 

Potentilla bipinnatifida 


Vol. 119 


Primula nutans 

Rosa woodsii 

Rumex maritimus ssp. fueginus 

Sagina nivalis 

Salix niphoclada 

Saxifraga nelsoniana ssp. pacifica 
Saxifraga rufopilosa 

Schoenoplectus acutus (Scirpus acutus) 
Schoenoplectus tabernemontani (Scirpus validus) 
Tricaphorum alpinum (Scirpus hudsonicus) 
Silene acaulis ssp. subacaulescens 

Sorbus sitchensis 

Stellaria umbellata 

Symphyorichum ciliatum (Aster brachyactis) 
Symphyorichum yukonense (Aster yukonensis) 
Taraxacum lyratum 

Typha latifolia 

Triantha glutinosa 

Utricularia minor 

Valeriana sitchensis 

Veronica americana 

Viola adunca 


Range Extensions of Introduced Taxa within the 
Yukon Territory (8): 

Alopecurus geniculatus 

Alopecurus pratensis 

Avena sativa 

Phalaris arundinacea 

Polygonum convolvulus 

Psathyrostachys juncea 

Secale cereale 

Vicia americana 


Comments on Native Taxa in the Yukon Flora (2): 
Lonicera involucrata 
Saxifraga nelsoniana ssp. carlottae 


Deletions of Native Taxa from the Yukon Flora (4): 
Clematis occidentalis 

Lonicera involucrata 

Polygonum persicaria 

Verbena hastata 


LYCOPODIACEAE 

Lycopodium sitchense Rupr. (L. sabinifolium Willd. 
ssp. sitchense (Rupr.) Calder & Taylor, Diphasiastrum 
sitchense (Rupr.) Holub), Sitka Club-moss — YUKON: 
Kluane National Park, Fisher Glacier near Alsek River, 
W-facing slope above valley glacier leading into Fish- 
er Glacier, 60°08'16"N 138°13'28"W, B. A. Bennett 03- 
983, 7 July 2003 (DAO); Kluane National Park, Alsek 
River, ca. 38 km SW of Haines Junction, 1245 m in al- 
pine Festuca altaica community, coarse soil, well 
drained aspect N62E slope 5% 60°28'N 137°52'W, 
G. W. Douglas & M. J. Ratcliffe 10779, 8 July 1978, on 
mixed sheet with Diphasiastrum alpinum, determined 
by B. A. Bennett, confirmed by A. Ceska. (Specimen 


housed at the Royal BC Museum V 137472). 

The map in Hultén (1968) had a dot in the southwest of 
the Yukon which Cody (1996) considered to be a misidenti- 
fication. Cody (1996) suggested that it should be looked for 
in that area and the specimens cited above is a verification. 


2005 


nae Y aca 
Saabs 


Kluane 


KLUANE 


= Logan NATIONAL PARK 


Warden Cabin 


s 
oF 
x 
S 
3 


WRANGELL - SAINT ELIAS 
NATIONAL PARK 


__GULF OF ALASKA 


Copy, BENNETT, AND CASWELL: VASCULAR PLANTS IN THE YUKON VII 


419 


\ 
\ 
& 
ty 


Aishihik > 
Q 


Lake \\ \ 

Laberge v 
—— ~ bg a j 
Haines Junction } \ 


Dezadeash 
Lake 


FIGURE |. Locator map — Southwest Yukon (courtesy of the Yukon Territorial Government). 


PTERIDACEAE 

Cryptogramma crispa (L.) R.Br. var. sitchensis (Rupr.) 
C. Christensen, Sitka Parsley Fern. —- YUKON: Kluane 
National Park, Fisher Glacier near Alsek River, W- 
facing slope above valley glacier leading into Fisher 
Glacier, mid-slope in large talus, thin soil of silty sand 
with rock outcrop, 60°08'16"N 138°13'28"W, B. A. 
Bennett 03-1000, 7 July 2003 (DAO); Fisher Glacier 
near Alsek River, NE of base camp, under boulders in 
steep unvegetated draws on south-facing slope, 
60°08'31"N 138°13'13"W, B. A. Bennett 03-1069, 8 
July 2003 (DAO). 

Douglas et al. (1981) did not include this fern in the Rare 
Vascular Plants of the Yukon but Cody (1996) knew it from 
only three sites adjacent to the Canol Road. The specimens 
cited above are an extension of the known range in the Ter- 
ritory of about 325 kilometers south of a site adjacent to the 
South Canol Road and are new to Kluane National Park. 


ASPIDIACEAE 

Athyrium filix-femina (L.) Roth var. cyclosorum 
(Ledeb.) Moore, Lady Fern — YUKON: moist organic 
soil, meadows in thickets, east Haines Hwy. at Km 152, 
60°02.970'N 136°53.001'W, P. Caswell 03-747, 21 
Aug. 2003 (DAO); Kluane National Park, Fisher Glac- 
ier near Alsek, NE of base camp, under boulders in 


steep unvegetated draws on south-facing slope where 
it was rare, 60°03'31"N 138°13'W, B. A. Bennett 03- 
1062, 8 July 2003 (DAO). 

This species was considered rare in the Territory by Dou- 
glas et al. (1981) and Cody (1996). The first specimen cited 
above is from a site about 60 kilometers east of a site mapped 
by Cody (1996) in southern Kluane National Park. The sec- 
ond specimen is from a site about 20 kilometers northwest. 


Polystichum lonchitis (L.) Roth., Holly Fern — YUKON: 
Kluane National Park, Fisher Glacier near Alsek River, 
NE of base camp, under boulders in steep unvegetat- 
ed draws on south-facing slope, where it was rare, 
60°08'31"N 138°13'13"W, B. A. Bennett 03-1061, 8 
July 2003 (DAO). 

This species was considered rare in the Yukon by Dou- 
glas et al. (1981) and Cody (1996) where it was only known 
from two widely separated localities. The specimen cited 
above is an extension of about 30 kilometers northwest of a 
previously known site in Kluane National Park adjacent to 
the British Columbia border. 


PINACEAE 

Pinus contorta Dougl. ex Loud. ssp. latifolia (Engelm.) 
Critchfield, Lodgepole Pine — YUKON: open slope west 
of the road, only single large old tree at least 10 m tall; 
appears to pre-date the construction of the highway, 


420 


appearing natural, Haines Road Km 157, south of 
Million Dollar Falls, 60°05'27"N 136°55'01", B. A. 
Bennett 03-003, 27 April 2003 (DAO). 

The specimen cited above is the southernmost yet found 
in the southwest of the Territory. It is from a site about 30 
kilometers south-southeast of a site reported by Cody et al. 
(2003) adjacent to the St. Elias Trail in Kluane National Park. 


TYPHACEAE 
Typha latifolia L., Common Cattail — YUKON: exten- 
sively placer mined area, Lower Hunker Creek area 
east of Dawson City, 64°01'N 139°03'01.2"W, G. Brun- 
ner 594, 7 Sept. 2003 (DAO); Gravel around and in 
small pond with scattered Salix, Carex, and Schoeno- 
plectus, up Hunker Creek Road east of Dawson Creek, 
64°O1'11.9"N 139°08'57.8"W, Peterson 18518, Saarela 
and Smith, 11 July 2004 (DAO). 

The specimen cited above is an extension of the known 
range of this rare species in the Territory of about 175 kilo- 
meters northwest of Mayo. 


POTAMOGETONACEAE 
Potamogeton praelongus Wulf., White-stemmed 
Pondweed — YUKON: floating on surface of lake, Pine 
Lake, 60°43.015'N 137°28.287'W, P. Caswell 03-491, 
23 July 2003 (DAO). 

Douglas et al. (1981) considered this taxon rare in the 
Territory. The specimen cited above is an extension of the 


known range of about 60 kilometers north of a site reported 
by Cody et al. (2000). 


NAJADACEAE 

Najas flexilis (Willd.) Rostk. & Schmidt, Wavy Water 
Nymph (Figure 2) — YUKON: meso-eutrophic lake and 
mesotrophic mire, west side of Wye Lake, Watson 
Lake, 60°07'N 128°48'W, R. Elven & H. Solstad 03- 
1301, 29 Aug. 2003 (DAO). 

This is a new species in the flora of the Yukon Territory 
which should be added to the list of rare plants (Douglas et 
al. 1981). Najadaceae can be separated from Potamogetona- 
ceae as follows: 

A. Flowers solitary in the leaf axils; 

leaf margins minutely serrulate with 

SAECO MEE ML DELISIGE sec, ahs ciel sois, tea dllnt cae Najadaceae 
A. Flowers born on spikes or racemes; 


leaf margins entire! oo oo. f Potamogetonaceae 


Najas flexilis can be described as follows: Submerged, annual 
monoecious, slender-stemmed, bushy or elongate, and often 
forming extensive beds; leaves numerous, becoming more 
crowded toward the branch tips, narrow, 1-3 cm long, mostly 
less than | mm wide, gradually tapering to a sharp point, 
minutely toothed; flowers unisexual, born in the axils of the 
branches and from the sheaths of leaf-bases; anthers 1|-celled; 
pistils mostly with 3 stigmas; fruit and seed narrowly ovoid 
or elliptic, 2-3 mm long; seed entirely filling the fruit, shiny, 
reddish-brown. 

In Canada, this species is known from Newfoundland to 
British Columbia; the nearest known site to that cited above 
is in Wood Buffalo Park just south of Fort Smith (Porsild and 
Cody 1980). 


POACEAE 
Agrostis humilis Vasey, (Agrostis thurberiana Hitchc., 
Podagrostis thurberiana (Hitchc.) Hultén), Alpine 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


1mm 


seed 


2mm 


FIGURE 2. Najas flexilis (Willd.) Rostk. & Schmidt, Wavy 
Water Nymph (illustration by J. R. Janish courtesy 
of the University of Washington Press). 


Bentgrass (Figure 3) — YUKON: Kluane National Park, 
rare in open herbaceous and graminoid meadows of 
south-facing mid-slopes in drier patches of open soil, 
Fisher Glacier near Alsek River, NE of base camp, 
60°03'31"N 138°13'13"W, B. A. Bennett 03-1073, 8 
July 2003, (DAO); Kluane National Park, Shursho 
Lake, meadow near lake, 100% vegetative coverage, 
moist organic soil, 60°03.6'N 137°23.8'W, P. Caswell 
& R. Maraj 04-427, 4 August 2004 (DAO) (deter- 
mined by S. J. Darbyshire). 

Agrostis humilis can be described as follows: Perennial 
tufted or matted grass from fibrous roots; stems 5-30 cm tall; 
leaves about 2.5 mm broad, panicle narrow; spikelets about 
2 mm long, violet; lemmas 1|.5-2.3 mm long, awnless; rachilla 
minutely pubescent; paleas present, at least half as long as 
the lemmas, anthers about (0.5) 06-0.8 mm long. 

This species is new to the flora of the Yukon Territory 
and should be added to the list of rare plants in the Territory. 
It is frequent in the subalpine and alpine zones throughout 
British Columbia, and extends into southern Alberta and 
south to California, Nevada, Utah and New Mexico. Agrostis 
gigantea and A. capillaris are rhizomatous; however, A. 
humilis is caespitose, so is more closely related to A. scabra 
and A. exarata. It can be separated from Agrostis scabra 
and Agrostis exarata as follows: 

A. Paleas present, at least half as long as lemmas 

“asa ou vsibb se lbasastte cc oe ie ik Elks wags eet ee A. humilis 
A. Paleas absent or less than half as long as lemmas 

Cah ae Mts eats ear ta PUM to te Dae A. scabra and A. exarata 


2005 


Alopecurus geniculatus L., Water Meadow-foxtail — 
YUKON: Fort Selkirk, 62°47'N 137°23'W, J. B. Tar- 
leton, Summer 1899 (US). 

This species, which is introduced in Canada from Eurasia, 
was reported as new to the Territory (Cody et al. 2003) from 
Horseshoe Slough. It is common in SW British Columbia 
north to the Queen Charlotte Islands. The above collection 
was reported by Britton and Rydberg (1901) and is a range 
extension of 130 km to the southwest. 


Alopecurus pratensis L., Meadow Foxtail — YUKON: 
uncommon in moist areas between fence and high- 
way, Takhini Salt Flats, 60°51'23"N 135°42'55"W, B. 
A. Bennett, R. Elven & H. Solstad 03-086, 26 Aug. 
2003 (B. A. Bennett Herbarium, photo DAO). 

This is an introduced species that is not yet common in 
the Territory. The nearest known site to that listed above is 
about 60 kilometers south of Whitehorse. 


Avena sativa L., Oats — YUKON: moist, gritty organic 
soil, Alaska Hwy., about 30 kilometers northwest of 
Haines Junction in squirrel research area, 60°57.110'N 
138°02.388'W, P. Caswell 03-563, 8 Aug. 2003 (DAO). 

Cody (1996) knew this introduced species from only 
three sites in the Territory (the vicinity of Mayo, North Canol 
Road and east of Watson Lake). An additional site was found 
by Cody in the vicinity of Whitehorse (Cody et al. 2004). 


Bromus japonicus Thunb. ex Murray, Japanese Brome 
(Figure 4) — YUKON: highway embankment, Alaska 
Hwy. at Jarvis River Bridge, 60°55.349'N 137°53.060'W, 
P. Caswell 03-555, 8 Aug. 2003 (DAO) (determined 
by S. J. Darbyshire). 

The specimen cited above is a new introduction to the 
Yukon Territory but is not known to persist. It could not be 
found at this location in 2004. It is known across Canada 
from Quebec to southern British Columbia from dry fields 
and waste places. 

Bromus japonicus can be described as follows: Annual, 
from fibrous roots. Culms (20-) 30-70 cm tall, erect or ascend- 
ing; sheaths mostly very densely pilose to lanate, the upper 
ones sometimes pubescent to glabrous; ligules hairy; sheaths 
densely haired; blades 10-20 cm long, mostly softly pilose 
on both sides; panicle 10-17 (-22) cm long, diffuse nodding, 
with spreading to ascending, to somewhat drooping, slender, 
flexuous branchlets that are mostly longer than the spikelets; 
spikelets 6-12 flowered, glumes glabrous to scabrous, lance- 
olate, the lower one 3-nerved 4-6 mm long, the upper 5- 
nerved 6-8.5 mm long; anthers included in the lemmas 1- 
1.5 mm long. It can be separated from Bromus hordeaceus 
as follows: 

A. Panicle dense or narrow, usually erect; 

panicle branchlets and pedicels shorter 

EREA ES INUCLEES 52,8 oars esl oy pe aise oe, cs B. hordeaceus 
A. Panicle nodding or erect, mostly open; 

panicle branches and pedicels as long as 

or longer than spikeletsi!04)) .../qesa diy: B. japonicus 


Festuca brachyphylla Schultes & Schultes f., Alpine 
Fescue — YUKON: Kluane National Park, nunatak be- 
tween Lowell and Dusty Glaciers near Ulu Mountain, 
60°21'32"N 138°34'49"W, B. A. Bennett 03-948, 6 
July 2003 (DAO) (determined by S. J. Darbyshire). 


Copy, BENNETT, AND CASWELL: VASCULAR PLANTS IN THE YUKON VII 


42] 


FIGURE 3. Agrostis humilis Vasey, Mountain Bent Grass 
(from Vasey, 1893. Grasses of the Pacific Slope) 


The specimen cited above is an extension of the known 
range in the Park of about 50 kilometers south of a site 
mapped by Cody (1996). 


Festuca brevissima Jurtz., Alaska Fescue — YUKON: 
Kluane National Park, loose rocks with some organic 
soil on rocky slope, Kings Throne Trail, 60°33.476'N 
139°14.282'W, P. Caswell 02-125, 21 June 2002 (DAO). 

The specimen cited above of this Amphi-Beringian spec- 
ies 1s an extension of the known range in the Territory of 
about 350 kilometers south of a site about latitude 63°40'N. 
It is new to the flora of Kluane National Park. 


Festuca minutiflora Rydb., Little Fescue — YUKON: 
Kluane National Park, loose soil with organic compo- 
nent at entrance of Arctic Ground Squirrel burrow, hill 
overlooking Ogilvie Glacier at termination of stream 
flowing south of Warden Cabin, nunatak at Logan 
Warden Cabin, 60°47.652'N 140°46.303'W, P. Caswell 
et al. 03-334, 12 July 2003 (DAO). 

The specimen cited above is the westernmost yet found 
in Kluane National Park and is an extension of the known 
range in the Park of about 100 kilometers to the west of a 
site mapped by Cody (1996). 


Leymus mollis (Trin.) Pilger ssp. mollis s.1., Ameri- 
can Lyme Grass — YUKON: roadside gravel, Takhini 
Hotsprings Road about | km east of hotsprings, one 


422 


patch only — introduced, 60°50'30"N 135°20'30"W, 
B. A. Bennett, R. Elven & H. Solstad 03-1300, 26 
Aug. 2003 (DAO). 

Cody (1996) knew ssp. villosissimus (Scribn.) A. Love 
from adjacent to the Arctic coast. The specimen cited above 
is the first record of this variable subspecies from the interi- 
or and southern part of the Territory. 


Phalaris arundinacea L., Reed Canary Grass — YUKON: 
3 km west of White River along Alaska Highway on 
north side of road; appears to be seeded, 61°59'N 
140°30'10"W, B. A. Bennett 04-0002, 10 April 2004 
(DAO). 

The specimen cited above is from a site about 225 kilo- 
meters northwest of a site southwest of Haines Junction in 
Kluane National Park cited by Cody et al. (2003) where it 
was native and a site about 175 kilometers to the northeast 


by Mechanic Creek where it was introduced (Cody et al. 
2001). 


Phippsia algida (Sol.) R.Br., Ice Grass — YUKON: 
Kluane National Park; dried and cracked mud at edge 
of large pond near Warden Cabin, nunatuk at Logan 
Warden Cabin, 60°47.628'N 140°45.148'W, P. Caswell 
et al. 03-216, 11 July 2003 (DAO). 

The specimen cited above, which is the most southwest- 
erly yet found in the Territory, is from a site about 115 kilo- 
meters southwest of a site mapped by Cody (1996) near the 
south end of Kluane Lake. 


Psathyrostachys juncea (Fisch.) Nevski (Elymus 
junceus Fisch.) Russian Wild Rye — YUKON: roadside 
at milepost 812, Alaska Highway, S. E. Welsh & G. 
Moore 7635, 2 July 1968 (University of Alaska Herb- 
arium, photo DAO) (determined by M. Barkworth). 
Stanley Welsh (1974) reported the specimen cited 
above and another one collected by M. V. Guttman 
collected from adjacent to Mile 1019 Alaska Hwy. on 
25 July 1960 (ISC). These records were unfortunate- 
ly missed when writing the Yukon Flora (Cody 1996, 
2000). 

This introduced species was first reported and described 
by Cody et al. (2003) sub. Elymus junceus from Dezadeash 
Lake adjacent to Km 195 Haines Hwy. 


Secale cereale L., Rye — YUKON: disturbed sandy 
soil, Whitehorse, between Mt. McIntyre Recreation 
Centre and Aquatic Centre Parking Lot, 60°03.32'N 
135°43.49'W, B. A. Bennett 03-1369, 8 Oct. 2003 (B. 
A. Bennett Herbarium, photo DAO). 

This introduced species was first collected by J. M. Gillett 
in gravel areas near warehouses in Watson Lake in 1949. 
The specimen cited above is only the second found growing 
wild in the Territory. 


CYPERACEAE 

Blysmopsis rufa (Huds.) Oteng- Yeboah (Scirpus rufus 
(Huds.) Schrad., Blysmus rufus (Huds.) Link), Red 
Bulrush, Swaying Rush (Figure 5) — YUKON: alkaline 
lake, saline flat 600 m south of Fox Creek, on east 
side of North Klondike Hwy., Km 227, 61°05.477'N 
135°16.826'W, R. Elven & H. Solstad 03-1305, 25 
Sept. 2003 (B. A. Bennett Herbarium, photo DAO). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


lodicules 


4 } 

HY, / 
/ 

¢ // 

hit 


\ 


od 


yea VER IS 


Bromus japonicus 


FIGURE 4. Bromus japonicus (Willd.) Rostk. & Schmidt, Wavy 
Water Nymph (Japanese Brome) (illustration by J. 
R. Janish courtesy of the University of Washington 
Press). 


2005 


This is a new species to the flora of the Yukon Territory 
which should be added to the list of rare plants (Douglas et 
al. 1981). To the east it is rare in the former District of 
Mackenzie from a saline meadow adjacent to the Keele River 
in the Mackenzie Mountains and on saline river banks near 
Wrigley in the Mackenzie Valley and to the west from tidal 
flats in the Matanuska area. It can be separated from Scho- 
enplectus tabernaemontani (Scirpus validus) as follows: 

A. Spikelets several, 2-ranked, in a terminal 
spike; perianth bristles lacking, 
immerescence Mattened ......).6. 5 se Blysmopsis rufa 
(Scirpus rufus) 
A. Spikelets solitary or few to numerous, 
not 2-ranked, in compact to open umbels 
or panicles; inflorescences not 
MAMeNEM. (fe 2. Schoenplectus tabernaemontani 
(Scirpus validus) 

Blysmopsis rufa (Scirpus rufus) can be described as: Peren- 
nial, loosely caespitose with horizontal rhizomes 1-3 mm thick; 
culms (5-) 10-45 cm x 1-2 (3) mm, glabrous; leaves 1-3; blades 
2-12 (-18) cm x 1.5-4 mm, midrib inconspicuous, apex blunt, 
glabrous; inflorescence in loose spikes, 5-8 spikelets distic- 
hous, spreading-ascending; bracts erect to oblique, leaflike or 
scalelike, 4-15 mm; perianth bristles separating from ach- 
ene when mature; style as long as stigma. 


Carex bebbii (L. H. Bailey) Olney ex Fern., Bebb’s 
Sedge (Figure 6) — YUKON: on old road to wetland 
west of bridge, north of road, La Biche River, 
60°05.74'N 124°01.98'W, B. A. Bennett 98-634, 15 
June 1998 (DAO) (determined by A. A. Reznicek). 
This species is new to the flora of the Yukon Territory 

and should be added to the list of rare species (Douglas et 
al. 1981). In the present key Carex bebbii would key out as 
C. crawfordii, but can be distinguished from that species 
using the following characters: 
A. Perigynia subulate to narrowly ovate- 

lanceolate, 3-4 times as long as wide, 

tinimand seale-like. ..b. sas. oe ess C. crawfordii 
A. Perigynia narrowly to broadly ovate, 

at most twice as long as wide, 

MIGHT AOS. le Ne de ote we Cees on. C. bebbii 


Carex brunnescens (Pers.) Poir., Brownish Sedge — 
YUKON: Kluane National Park, very wet, muddy soil 
at base of beaver dam, trail between Mush Lake and 
Bates Lake, 60°18.047'N 137°31.560'W, P. Caswell 
03-699, 30 July 2003 (DAO). 

The specimen cited above is an extension of the known 
range in the Park of about 40 kilometers south of a site at 
60°42'N 137°27'W reported by Cody et al. (2003). These 
two sites are about 200 kilometers west of a site adjacent to 
Jake’s Corner (Cody 1996). 


Carex buxbaumii Wahlenb., Buxbaum’s Sedge — 
YUKON: bog with tussocks at edge of Haines Hwy., 
60°08.570'N 136°58.469'W, P. Caswell 02-454, 13 
July 2002 (DAO); Kluane National Park, bank of shady 
streamlet shaded by willows, Auriol Trail, 64°44'15"N 
137°30'40"W, P. Caswell 02-662, 7 Aug. 2002 (DAO); 
growing in edge of small thermokarst pond south- 
west of highway, Alaska Hwy., 6 km northwest of 
Dry Creek #2, 62°13.455'N 140°41.544'W, B. A. Ben- 
nett, R. Elven & H. Solstad 03-087, 8 Aug. 2003 (DAO). 


Copy, BENNETT, AND CASWELL: VASCULAR PLANTS IN THE YUKON VII 


423 


Fa? ) PH VOW, Bao 
SU, 
‘ N 


N 


FIGURE 5. Blysmopsis rufa (Huds.) Oteng- Yeboah, Red Bul- 
rush (drawn by Lee Mennell). 


424 


This species was not included in the rare plants of the 
Territory (Douglas et al. 1981) because it is widespread cir- 
cumpolar. The first specimen cited above is from a site about 
125 kilometers southwest of Whitehorse. The second speci- 
men is from a site just southwest of Haines Junction and is 
the second record for Kluane National Park. The third spec- 
imen is an extension of about 120 kilometers northwest of a 
site north of Kluane Lake. 


Carex canescens L., Grey Sedge — YUKON: Kluane 
National Park, Cyclops Lake, about 10 km NE of Mush 
Lake, 60°21'20"N 137°11'30"W, R. D. Wickstrom 276, 
21 July 1974 (DAO); gravel with some organic soil, 
base of highway embankment, streamlet west of Alas- 
ka Hwy. near Rock Glacier, 60°25.0'N 137°02.5'W, 
P. Caswell 02-654, 5 Aug. 2002 (DAO). 

This species is widespread throughout much of the Yukon 
Territory. The specimens cited above are new to Kluane 
National Park. The nearest site known to Cody (1996) is 
south of Whitehorse, about 125 kilometers to the east. 


Carex eburnea Boott, Bristle-leaved Sedge — YUKON: 
growing in crevice of rockface, steep scree slope, old 
burn, Beaver River, 60°24'44"N 125°48'0S"W, B. A. 
Bennett 97-456, 14 Aug. 1997 (DAO); dry scree below 
outcrop, Beaver River Camp #1, 60°30'40"N 
126°06'26"W, B. A. Bennett 97-417, 13 Aug. 1997 
(DAO); raven nest site, riverbar, base of cliff, Wind 
River, 65°12.49'N 135°13.17'W, B. A. Bennett 00- 
369, 5 July 2000 (DAO); silty seep on edge of river, 
Peel River Site #1, 65°58.06'N 134°49.69'W, B. A. 
Bennett 00-433, 10 July 2000 (DAO) (determined by 
A. A. Reznicek); in floodplain tall closed balsam 
poplar, White River, 63°07.01'N 140°23.27'W, R. Rosie 
s.n., 26 Aug. 2002 (B. A. Bennett Herbarium, photo 
DAO); Dempster Highway, Windy Pass, Km 160, flat 
floodplain with Leymus innovatus in sandy calcareous 
soul, Bennett & Mulder 03-1307, 30 Aug. 2003 (DAO). 
Douglas et al. (1981) knew this rare species in the Terri- 
tory from a single collection in the Koidern area in the 
extreme west just south of latitude 62°N. The first four speci- 
mens cited above which were originally determined as Carex 
capillaris and C. williamsii extend the known range to the 
extreme southeast and northeast to the Wind and Peel rivers. 
The additional two specimens extend the known range 
westward to the Dempster Highway and White River. 


Carex lasiocarpa Ehrh., Slender Sedge — YUKON: 
flooded shoreline, Blind Lake, 60°05'0.5"N_ 128° 
14'49.7"W, J. Staniforth 6, 26-28 June 2003 (DAO) 
(determined by P. M. Catling); small pond near high- 
way northwest of bridge, mineralized soil, northwest 
side of Donjek River, 61°40.8'N 139°45.58'W, B. A. 
Bennett, R. Elven & H. Solstad 03-056, 8 Aug. 2003 
(DAO). 

This species was considered rare in the Territory (Douglas 
et al. 1981) on the basis of a collection from the Elsa area. 
An additional site was mapped in the southwest by Cody 
(1996). Cody et al. (1998, 2002) reported new sites in the 
southeast from the vicinities of Watson Lake and Frances 
Lake. The first specimen cited above is an extension of the 
known range of about 30 kilometers east of Watson Lake. 
The second specimen is an extension of the known range of 


THE CANADIAN FIELD-NATURALIST 


Volit9 


FIGURE 6. Carex bebbii (L.H. Bailey) Olney ex Fern., Bebb’s 
Sedge (illustration by J. R. Janish courtesy of the 
University of Washington Press). 


2005 


about 60 kilometers north of a site in northern Kluane Nation- 
al Park. 


Carex laxa Wahlenb., Weak Sedge — YUKON: edge of 
small thermokarst pond southwest of Alaska Highway 
6 kilometers northwest of Dry Creek, 62°13.455'N 
140°41.544'W, B. A. Bennett, R. Elven & H. Solstad 
03-042, 8 Aug. 2003 (Yukon Government Herbarium, 
photo DAO). 

This Amphi-Beringian species was previously known in 
North America from two stations in Alaska, one in the 
Mackenzie Delta, and two localities in Central Yukon where 
it was considered rare by Douglas et al. (1981) [Cody 1996]. 
The specimen cited above is from a site about 300 kilome- 
ters southwest of sites in the vicinities of Mayo and Elsa. 


Carex lenticularis Michaux var. dolia (M. E. Jones) 
L. A. Standley (C. enanderi Hultén), Enander’s Sedge 
— YUKON: Kluane National Park, rare, seen only along 
creek near camp, Fisher Glacier near Alsek River, 
base camp, 60°08'31"N 138°13'13"W, B. A. Bennett 
03-955, 6 July 2003 (DAO) (determined by Ph.D. can- 
didate Julie A. Dragon from the University of Vermont). 

First reported from Yukon by Standley et al. (2002) this 
is a new variety in the Yukon Territory and should be added 
to the Rare Vascular Plants of the Yukon (Douglas et al. 1981). 
It can be separated from Carex lenticularis var. lenticularis 
as follows (from Flora of North America Volume 23): 

A. Terminal spike gynecandrous (with 

pistillate flowers at the apex); 

proximal spike not exceeding 1.5 cm; 

peduncle less than | cm; perigynia 

COMO GH de PE Gs RU SS NG ae ae a var. dolia 
A. Terminal spike usually staminate; 

proximal spike longer than 1.5 cm; 

peduncle | cm or longer; perigynia 

Ovord toellipsoid  / 5.42 i eee sos var. lenticularis 

Carex eleusinoides is similar in inflorescence morpholo- 
gy, but may be distinguished by its red basal sheaths and 
short, veinless perigynia. 

This species was considered rare in the Territory by Dou- 
glas et al. (1981). The specimen cited above, which is very 
reduced in size, is new to Kluane National Park and is from 
a site about 140 kilometers south of a site at the north end 
of Kluane Lake. 


Carex lenticularis Michx. var. lipocarpa (Holm) L. A. 
Standley, Kellogg’s Sedge — YUKON: Kluane Nation- 
al Park, braided river bed in muddy area toward a 
small pond, Plug Creek, 60°03.593'N 138°12.055'W, 
P. Caswell et al. 03-672, 29 July 2003 (DAO). 

Cody (1996) suggested that this sedge should be looked 
for in southern Yukon Territory because Hultén (1968) indi- 
cated collection sites from just north of where the British 
Columbia specimens were found. Bruce A. Bennett found it 
in a Sphagnum bog southeast of Watson Lake in 1996 (Cody 
et al. 1998). This record was overlooked by Standley et al. 
(2002). The specimen cited above is only the second yet con- 
firmed and is from a site about 500 kilometers to the west. 


Carex livida (Wahl.) Willd., Pale Sedge — YUKON: 
southwest of highway, growing in edge of small ther- 


mokarst pond, Alaska Hwy., 6 km NW of Dry Creek 
#2, 62°13.455'N 140°41.555'W, B. A. Bennett 03-043, 


Copy, BENNETT, AND CASWELL: VASCULAR PLANTS IN THE YUKON VII 


425 


1 mm 


FIGURE 7. Carex nigricans C.A. Mey., Blackish Sedge. (illus- 
tration by J. R. Janish courtesy of the University of 
Washington Press). 


8 Aug. 2003 (DAO); moist organic soil, bog east of 
Haines Hwy., 60°08.421'N 136°58.499'W, P. Caswell 
02-448, 13 July 2002 (DAO). 

This circumpolar species with large gaps in its range was 
considered rare in the Territory by Douglas et al. (1981). The 
first specimen cited above is an extension of the known range 
in the Territory of about 160 kilometers west of a site south- 
west of Carmacks. The second specimen is from a site about 
200 kilometers south of the site southwest of Carmacks. 


Carex microglochin Wahl., Few-seeded Fen Sedge — 
YUKON: moist organic soil in bog east of Haines Hwy., 
60°08.421'N 136°59.499'W, P. Caswell 02-452B, 13 
July 2002 (DAO). 

This circumpolar species was known to Cody (1996) from 
four widely separated areas in the Territory. The specimen 
cited above is from a site about 85 kilometers southeast of a 
site in Kluane National Park just southwest of Haines Junction. 


Carex microptera Mack., Small-winged Sedge — 
YUKON: wet lakeshore, Inlet of Caribou Creek to Cari- 
bou Lake, 60°32.0'N 134°15.9'W, S. Withers SWOO- 
168, 7 Aug. 2000 (DAO). 

This collection originally identified as Carex pachystachya 
(Cody et al. 2003) was reidentified by A. A. Reznicek. Dou- 
glas et al. (1981) did not consider this species rare in the 


426 


Territory because it is widespread in western North Ameri- 
ca. Cody (1996) knew it from only nine sites in the southern 
part of the Territory. The specimen cited above is from a site 
intermediate between the vicinities of Whitehorse and John- 
son’s Crossing. 


Carex nardina Fries, Spikehead Sedge — YUKON: 
Kluane National Park, nunatak near Ulu Mountain, 
60°21'32"N 138°34'49"W, B. A. Bennett 03-929, 6 
July 2003 (DAO). 

The specimen cited above is an extension of the known 
range in the Park of about 35 kilometers southwest of the 
most southeasterly site in the Park mapped by Cody (1996). 


Carex nigricans C. A. Mey., Blackish Sedge (Figure 
7) — YUKON: Kluane National Park, uncommon, seen 
along creek near camp and muddy flats upstream, 
Fisher Glacier near Alsek River, base camp, 60°08'31'"N 
138°13'13"W, B. A. Bennett 03-954, 6 July 2003 (DAO); 
300 m east of camp, flat moist tundra at bottom of broad 
valley with Diphasiastrum alpinum, Juncus drum- 
mondii, Fisher Glacier near Alsek River, 60°08'31"N 
138°13'13"W, B. A. Bennett 03-1074, 8 July 2003 
(DAO). 

This is a new species in the Yukon Territory and should 
be added to The Rare Vascular Plants of the Yukon (Douglas 
et al. 1981). Carex nigricans can be described as: plants 
loosely caespitose, mat-forming; rhizomes short, stout; culms 
5-30 cm; leaves flat to the tip, 2-4 mm wide; pistillate scales 
reddish brown to black, lanceolate, as broad as long and as 
long as or slightly longer than perigynia, margins not hya- 
line or scarious, apex acute to acuminate; perigynia 3.8-4.1 
x 1-1.2 mm; beak dark brown to black, frequently as long 
as body of perigynium; stigmas 3. It can be separated from 
Carex pyrenaica as follows: 

A. Plants rhizomatous; leaves 1.5-2 mm 

broad, flat or channelled basally; 

staminate flowers occupying one-third 

to"one=half of thespike: <0 20: 025 285. C. nigricans 
A. Plants clump-forming; leaves 1.5 mm 

broad or less, channelled throughout; 

staminate flowers occupying less than 

one-third of the Spiker) Sein May ee C. pyrenaica 


Carex parryana Dewey, Parry’s Sedge — YUKON: silty 
broad alkaline river flat, northwest side of Donjek 
River, 61°40.8'N 139°45.58'W, B. A. Bennett, R. Elven 
& H. Solstad 03-054, 8 Aug. 2003 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 125 kilometers 
northwest of a site in the vicinity of Kluane Lake. 


Carex phaeocephala Piper, Dunhead Sedge — YUKON: 
Kluane National Park, west-facing slope, drier soils 
in herbaceous meadow dominated by Luetkea pecti- 
nata, Fisher Glacier near Alsek River, 60°08'09"N 
138°13'10"W, B. A. Bennett 03-986, 7 July 2003 (DAO); 
common open herbaceous and graminoid meadow of 
south-facing mid-slopes in drier patches of open soil, 
northeast of base camp, Fisher Glacier near Alsek 
River, 60°08'31"N 138°13'13"W, B. A. Bennett 03- 
1065, 8 July 2003 (DAO) (determined by A. A. Rez- 
nicek); south-facing slope, thin coarse soil surround- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


ed by boulders and ice, nunatak between Lowell 
and Dusty glaciers near Ulu Mountain, 60°21'32"N 
138°34'49"W, B. A. Bennett 03-930, 6 July 2003 
(DAO) (determined by A. A. Reznicek). 

Douglas et al. (1981) knew this rare species in the Terri- 
tory only from Kluane National Park. The first two speci- 
mens cited above are an extension of about 70 kilometers 
west of a site adjacent to the Haines Hwy. mapped by Cody 
(1996) and the third specimen is about 40 kilometers further 
to the northwest. 


Carex sartwellii Dewey, Sartwell’s Sedge (Figure 8) 
— YUKON: occasional in alkaline area in grassy open- 
ings on dwarf birch-willow flats at 2500', Mile 26 on 
road from Whitehorse to Dawson, 61°08'N 135°21'W, 
J. A. Calder and I. Kukkonen 28035, 9 Aug. 1960 
(DAO) (determined by A. A. Reznicek). 

This is a new species to be found in the Yukon Territory 
and should be added to the list of rare species (Douglas et 
al. 1981). It can be separated from Carex praegracilis as 
follows: 

A. Upper sheaths translucent below; 

perigynia with beaks not sharply 

bidentate, not thin — margined 

ABOVE eB s OUT Oe? Ae C. praegracilis 
A. Upper sheaths green-lined below; 

perigynia with beaks sharply 

bidentate, thin margined above ........ C. sartwellii 

In Canada this species is known from south-central and 
southeastern British Columbia north to southern part of the 
former District of Mackenzie and east to Quebec. Carex 
sartwellii can be described as follows: Perennial herb from 
stout, creeping, scaly, fibre-covered rhizomes; stems 30-80 cm 
tall arising singly or a few together, longer than the leaves; 
sheaths tight, brown, glabrous, inner band green; blades 2-4 
per stem 2.5-5 mm wide, lower ones reduced to scales; spikes 
20 or more nearly cylindrical, except near the apex; basal 
spikes more prominent than middle spikes; perigynia (2.3-) 
2.8-4 x 1.3-2 mm; achenes brown, smooth 1.2-2 mm long, 
stigmas 2. 


Carex siccata Dewey, Hay Sedge — YUKON: Picea 
mariana dominated forest, Beaver River near Larsen 
Airstrip, 60°10'42"N 125°05'38"W, B. A. Bennett 
95314, 22 June 1995 (B. A. Bennett Herbarium, photo 
DAO) (determined by A. A. Reznicek). 

The specimen cited above was originally identified as 
Carex foenea (Cody et al. 1998) and all of the three speci- 
mens mapped previously (Cody 1996) are presumably 
referable to this species. The name Carex foenea correctly 
refers to a species 1n section Ovales (C. aenea), but has fre- 
quently been misapplied to this species. Carex foenea (= C. 
aenea of Cody 1996) is widespread in southern Yukon. The 
name Carex siccata was first applied to Yukon plants by 
Reznicek (2002). Carex foenea and C. siccata can be sepa- 
rated as follows: 

A. Plants caespitose; spikes 3-7 (-11); 
perigynia with wing 0.2-0.4 mm wide 

A. Plant with long creeping rhizomes; 
spikes 4-12; perigynia wingless ...... Carex siccata 


... Carex foenea 


Eriophorum gracile Koch, Slender Cotton-grass — 
YUKON: flooded shoreline, Blind Lake, 60°05'0.5"N 
128°14'49.7"W, J. Staniforth 3, 26-28 June 2003 (DAO). 


2005 


Cody (1994) reported the first specimen known from the 
Territory from a site near the British Columbia border between 
longitudes 127°W and 128°W and recommended that this 
species should be added to the list of rare vascular plants of 
the Yukon Territory (Douglas et al. 1981). Additional speci- 
mens have since been reported from Upper Coal River and 
Frances Lake (Cody et al. 1998) and Turner Lake (Cody et 
al. 2001). The specimen cited above is from a site about half 
way between the first known site and Watson Lake. 


Schoenoplectus acutus (Muhl. ex Bigelow) A. & D. 
Love var. acutus) (Scirpus acutus Muhl. ex Bigelow), 
Hard-stemmed Bulrush — YUKON: aquatic emergent, 
Blind Lake, 60°06'7.9"N 128°13'1.6"W, J. Staniforth 
7, 26-28 June 2003 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 300 kilometers southeast of 
sites in the vicinity of Ross River (Cody et al. 2001). 


Schoenoplectus tabernaemontani (C.C. Gmel.) Palla 
(Scirpus validus Vahl), Soft-stem Club-rush — YUKON: 
extensively placer mined area, Lower Hunker Creek 
area east of Dawson City, 64°01'14.2"N 139°09'01.2"W, 
G. Brunner 593, 7 Sept. 2003 (DAO). 

The specimen cited above is from a site about 150 kilo- 
meters south of a site adjacent to the Dempster Hwy. and 
about 175 kilometers northwest of a site in the vicinity of 
Mayo. 


Trichophorum alpinum (L.) Pers. (Scirpus hudsoni- 
anus (Michx.) Fern.), Hudson Bay Clubrush — YUKON: 
growing in edge of small thermokarst pond south- 
west of Alaska Hwy., 6 km NW of Dry Creek #2, 
62°13.455'N 140°41.544'W, B. A. Bennett, R. Elven 
& H. Solstad 03-044, 8 Aug. 2003 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 50 kilometers to the north- 
west of a site adjacent to the Alaska Hwy. mapped by Cody 
(1996). 


JUNCACEAE 

Juncus bufonius L. s.1., Toad Rush — YUKON: in shal- 
lows of northeast shore with Callitriche and Eleocharis, 
Hidden Lakes, Whitehorse, 60°41.9'N 135°00.6'W, 
B. A. Bennett 03-070, 13 Aug. 2003 (DAO); Kluane 
National Park, packed moist sand with small rocks and 
organic component, strip in middle of road, Alsek Trail, 
60°46.042'N 137°44.657'W, P. Caswell 03-755, 25 
Aug. 2003 (DAO). 

The first specimen cited above is from a site in the south- 
ern part of the Territory between sites in the vicinities of 
Haines Junction and east of South Canol Road. The second 
specimen is an extension of the known range in the Territory 
of about 250 kilometers southwest of a site south of Mayo 
mapped by Cody (1996). 


Juncus filiformis L., Thread Rush — YUKON: Kluane 
National Park, wet, muddy soil, small marsh at lake- 
shore, west end of Mush Lake, 60°18.377'N 137° 
32.68 1'W, P. Caswell 03-687, 30 July 2003 (DAO). 
The specimen cited above is an extension of the known 
range in the Territory of about 85 kilometers southwest of a 
site adjacent to the Alaska Hwy. east of Haines Junction 


Copy, BENNETT, AND CASWELL: VASCULAR PLANTS IN THE YUKON VII 


427 


Carex soartwellii 


FIGURE 8. Carex sartwellii Dewey, Sartwell’s Sedge. (illus- 
tration by J. R. Janish courtesy of the University of 
Washington Press). 


reported by Cody et al. (2004). It is new to Kluane National 
Park. 


Luzula piperi (Cov.) M. E. Jones, Piper’s Wood-rush 
— YUKON: Kluane National Park, uncommon in herba- 
ceous meadows, usually on the toe of slopes, Fisher 
Glacier near Alsek River, base camp, 60°08'31"N 
138°13'13"W, B. A. Bennett 03-972, 6 July 2003 
(DAO). 

The specimen cited above is only the third known to 
Cody (1996) in Kluane National Park. It is an extension of 
the known range to the southwest of about 40 kilometers. 


Luzula spicata (L.) DC., Spiked Wood-rush — YUKON: 
Kluane National Park, W-facing slope above valley 


428 


glacier leading into Fisher Glacier near Alsek River, 
60°08'25"N 138°13'20"W, B. A. Bennett 03-1016, 7 
July 2003 (DAO). . 

The specimen cited above is the southernmost yet found 
in the Park. It is about 40 kilometers south-southwest of a 
site southwest of Haines Junction. 


LILIACEAE 

Triantha glutinosa (Michx.) Baker (Tofieldia gluti- 
nosa ssp. brevistyla sensu Cody (1996)), Sticky False 
Asphodel — YUKON: silty broad alkaline river on flat, 
seasonally flooding shore, dominated by Salix brachy- 
carpa, Carex parryana, Calamagrostis stricta and Aster 
yukonensis, northwest side of Donjek River, 61°40.8'N 
139°45.58'W, B. A. Bennett, R. Elven & H. Solstad 
03-050, 8 Aug. 2003 (DAO). 

Douglas et al. (1981) considered this taxon as rare in the 
Territory. The specimen cited above is an extension of the 
known range of about 230 kilometers northwest of a site in 
the vicinity of Whitehorse where it was collected by C. E. 
Kennedy et al. in 1996. The scientific name Triantha gluti- 
nosa was brought forward by Douglas et al. (2001) and 
Utech (2002). 


IRIDACIAE 
Tris setosa Pall. ssp. interior (Anders.) Hultén — YUKON: 
southwest of highway, growing in edge of small ther- 
mokarst pond, Alaska Hwy. 6 km NW of Dry Creek 
#2, 62°13.455'N 140°41.544'W, B. A. Bennett, R. 
Elven & H. Solstad 03-048, 8 Aug. 2003 (B. A. Ben- 
nett Herbarium, photo DAO). 

This species was considered rare in the Territory by Dou- 
glas et al. (1981). The specimen cited above is from a site 
about 20 kilometers southeast of the Alaska border. 


ORCHIDACEAE 

Coeloglossum viride (L.) Hartm. ssp. bracteatum 
(Muhl. ex Willd.) Hultén, Bracted Green Orchid — 
YUKON: moist, black organic soil, edge of road shad- 
ed by poplar woods, old road west of Alaska Hwy. 
(between Kluane Wilderness Camp (Scully’s bar) and 
Donjek River), 61°36.701'N 139°37.730'W, P. Caswell 
02-189, 25 June 2002 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory on the basis of a single collection in the extreme 
south in the vicinity of Little Atlin Lake. Cody (1996) added 
a second site from adjacent to the Bonnet Plume River at 
latitude 65°10'N, and Cody et al. (2001/2002) added two 
additional sites east and west of the Bonnet Plume River 
site and a site near the Donjek River. The specimen cited 
above is an extension of the known range in the Territory of 
about 13 kilometers southeast of the Donjek River site. 


SALICACEAE 
Salix niphoclada Rydb. (S. brachycarpa Nutt. ssp. 
niphoclada (Rydb.) Argus, Barren-ground Willow — 
YUKON: Kluane National Park, Fisher Glacier near 
Alsek River, SW-facing above glacier, 60°08'06"N 
138°12'42"W, B. A. Bennett 03-1027, 7 July 2003 
(DAO) (determined by G. W. Argus). 

The specimen cited above is the southernmost yet found 
in Kluane National Park. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


SANTALACEAE 

Comandra umbellata (L.) Nutt. ssp. pallida (A.DC.) 
Piehl, Pale Commandra — YUKON: Kluane National 
Park, open, sage-covered sweeping slope overlook- 
ing Slims West Trail and west of the Sheep Bullion 
Trail, :60°59.138'N 138°35.832'W, BP Caswell é& ©. 
Freese 03-753, 3 Aug. 2003 (DAO). 

The specimen cited above is the first known of this species 
from Kluane National Park. It was however known to Cody 
(1996) from adjacent to Kluane Lake on the east side of the 
Alaska Hwy. Cody (1996) also mapped a site near the Five 
Finger Rapids adjacent to the Klondike Hwy. southeast of 
Fort Selkirk (F. Anderson, 18 June 1898, Hultén 1940). It has 
not been seen again in that area despite some searching. 
Hultén (1968) and Douglas et al. (1981) accidentally mapped 
it from the vicinity of Dawson. 


POLYGONACEAE 

Polygonum achoreum Blake, Striate Knotweed — 
YUKON: front yard of cabin, Lee’s Camp, Beaver River, 
60°11'S7"N 125°09'42"W, B. A. Bennett 97-493, 18 
Aug. 1997 (DAO) (determined by M. Costea). 

The specimen cited above, originally identified as P. bux- 
iforme (Cody et al. 2000), is an extension of the known 
range in the Territory of about 575 kilometers east of the 
vicinity of Whitehorse. 


Polygonum convolvulus L., Bind-weed, Wild Buck- 
wheat — YUKON: dry, very rich organic soil, abandoned 
vegetable garden, 14 Willow Acres, Haines Junction, 
60°45.823'N 137°29.458'W, P. Caswell 03-427, 17 
July 2003 (DAO). 

Cody (1996) knew this introduced species in the Territory 
from only four widely separated sites (Whitehorse, Dawson, 
Carmacks and adjacent to the Campbell Highway northwest 
of Ross River). 


Polygonum lapathifolium L. (P. pensylvanicum L. 
ssp. oneillii (Brenckle) Hultén — YUKON: usually sub- 
merged in shallow water in marly mud, Ear Lake, 
Whitehorse, 60°40.81'N 135°02.54'W, B. A. Bennett 
03-035, 9 Aug. 2003 (DAO); in shallows of northeast 
shore with Callitriche verna and Eleocharis acicu- 
laris, Hidden Lake, Whitehorse, 60°41.9'N 135°00.6'W, 
B. A. Bennett 03-071, 13 Aug. 2003 (DAO). 

Cody et al. (2000) reported this species, which was con- 
sidered rare in the Territory by Douglas et al. (1981), from 
three sites about 100 kilometers northwest of the specimens 
cited above. 


Polygonum persicaria L., Lady’s-thumb — Cody (1996, 
2000) reported the occurrence of this species based 
on a collection from the Canol Road Mile 55-60 on 
the east shore of Quiet Lake near Old Road Camp by 
A. E. Porsild & A. J. Breitung 11124, 4 July 1944 
(CAN). 


This specimen was reviewed by R. Staniforth (1975) 
who reported that it was possibly P. lapathifolium. It 
was reviewed again by L. L. Consaul (1987) who 
revised it to P. lapathifolium. P. persicaria should be 
deleted from the flora of the Yukon Territory. 


2005 


Rumex maritimus L. ssp. fueginus (Phil.) Hultén, 
Golden Dock — YUKON: dry, very rich organic soil, 
abandoned vegetable garden at 14 Willow Acres, 
Haines Junction, 60°45.823'N 137°29.458'W, P. 
Caswell 03-437, 17 July 2003 (DAO) (determined by 
S. J. Darbyshire). 

The specimen cited above is an extension of about 35 
kilometers southeast of a site adjacent to the Alaska Hwy. 
mapped by Cody (1996). 


CARYOPHYLLACEAE 
Minuartia dawsonensis (Britt.) House, Bog Sandwort 
— YUKON: Kluane National Park, packed earth on 
trail, Alsek Trail past Serpentine Creek, 60°41.397'N 
137°46.603'W, P. Caswell 02-220, 26 June 2002 (DAO). 
The specimen cited above is the first reported from Klu- 
ane National Park. It is from a site about 30 kilometers 
southeast of a site mapped by Cody (1996) adjacent to the 
Alaska Hwy. 


Sagina nivalis (Lindbl.) Fries — YUKON: Kluane 
National Park, frost heaved mud beside large rocks at 
edge of lake, shore of pond, south side of large pond 
in front of Warden Cabin, nunatak at Logan Warden 
Cabin, 60°47.628'N 140°45.148'W, P. Caswell et al. 
03-356, 3 July 2003 (DAO). 

The specimen cited above is only the second record of 
the occurrence of this rare species in Kluane National Park 
where it is about 175 kilometers northwest of the first site 
(Cody 1996). Douglas et al. (1981) considered this species 
rare in the Yukon Territory. 


Silene acaulis (L.) Jacq. ssp. subacaulescens (E.N. 
Williams) Hultén — YUKON: Kluane National Park, 
Fisher Glacier near Alsek River, base camp, 60°08'31"N 
138°13'13"W, B. A. Bennett 03-965, 6 July 2003 (DAO). 

The specimen cited above is only the second known from 
the Park. To the north the nearest sites are just south of lati- 
tude 64°N and to the east just east of longitude 133°W 
(Cody 1996). 


Stellaria umbellata Turcz., Umbellate Starwort — 
YUKON: Buckland Hills, Ivvavik National Park, 
69°25'N 139°38'W, R. Elven 7229-99, 5 Aug. 1999 
(University of Alaska Herbarium, photo DAO) (deter- 
mined by R. Elven). 

This species was considered rare in the Territory by Dou- 
glas et al. (1981). Cody (1996) knew it from only three widely 
separated sites. The specimen cited above is an extension of 
the known range of about 60 kilometers to the northwest of 
a site adjacent to the Babbage River. 


NYMPHAEACEAE 

Nuphar variegatum Engelm., Bullhead Lily — YUKON: 
pond south of the Alaska Hwy. and west of Donjek 
River, 61°41'11.8"N 139°47'01.6"W, D. Normandeau 
03-533, 26 July 2003 (DAO). 

This is a rare species in the Territory (Douglas et al. 
1981). The specimen cited above is an extension of the known 
range of about 230 kilometers southwest of a site south of 
Mayo mapped by Cody (1996). 


Copy, BENNETT, AND CASWELL: VASCULAR PLANTS IN THE YUKON VII 


429 


RANUNCULACEAE 

Clematis occidentalis (Hornem.) DC. ssp. grosseser- 
rata (Rydb.) Taylor & MacBryde, Purple Clematis — 
YUKON: Haines Junction, L. Fournier s.n., 13 July 1958 
(QFA). 

The sample of Clematis was revised by B. Boivin in 
1965 to C. tangutica, with the note: “Note many folioles, 
more likely to be C. tangutica, maybe cultivated.” Clematis 
tangutica is a species that has escaped from cultivation and 
persists in the vicinity of Whitehorse and Carcross (Cody et 
al. 2001). Clematis occidentalis should be deleted from the 
flora of the Yukon Territory. 


PAPAVERACEAE 
Papaver radicatum Rottb. ssp. kluanensis (D. Love) 
D. F. Murray — YUKON: alpine, Friday Creek, 60°25'N 
135°14 W, A. Rhodes 99-537, 11 Aug. 1999 (B. A. 
Bennett Herbarium, photo DAO) (determined by R. 
Elvin and H. Solstad). 

The specimen cited above is an extension of about 120 


kilometers east of the type site in Kluane National Park 
(Murray 1995). 


BRASSICACEAE 

Aphragmus eschscholtzianus Andrz., Eschscholtz’ Lit- 
tle Nightmare, Aleutian Cress — YUKON: Kluane 
National Park, moist, frost sorted rocks and gravel, 
Wade Mountain, 61°18.594'N 139°30.921'W, P. Cas- 
well 02-353, 7 July 2002 (DAO) (determined by G. 
A. Mulligan). 

Douglas et al. (1981) knew this rare species in the Terri- 
tory from only four sites in the southwest, three of which 
were in Kluane National Park. The specimen cited above is 
the fourth in the Park and is from a site about 45 kilometers 
east of the westernmost site mapped by Cody (1996). 


Arabidopsis salsuginea (Pallas) N. Busch (Thellungiel- 
la salsuginea (Pallas) O. E. Schulz), Saltwater Cress 
— YUKON: very rich organic soil, abandoned Llama pas- 
ture, Haines Junction, 14 Willow Circle, 60°45.816'N 
137°29.466'W, P. Caswell 03-414, 17 July 2003; same 
area, 60°45.843'N 137°24.484'W, P. Caswell 03-055, 
19 June 2003 (DAO) (determined by G. A. Mulligan). 

Douglas et al. (1981) and Cody (1996) knew this species 
from only five sites in the southwest of the Territory. The 
sixth site cited above is from a location just southeast of the 
westernmost previously recorded. 


Arabis boivinii G. A. Mulligan — YUKON: Kluane 
National Park, dry sand with organic component, Mush 
Lake Warden Cabin at west end of Mush Lake, 
60°18.496'N 137°32.776'W, P. Caswell 03-708, 31 
July 2003 (DAO) (determined by G. A. Mulligan). 

Cody et al. (2001) reported the first two known sites in 
the Territory from adjacent to the Haines Hwy. Cody et al. 
(2003) reported a third site from the old Experimental Farm 
northwest of Haines Junction. The specimen cited above is 
the first known from within the borders of Kluane National 
Park. 


Draba albertina Greene, Slender Draba, Alaska Whit- 
low-grass — YUKON: Kluane National Park, roadside 


430 


spruce forest, Alaska Hwy. at squirrel research area, 
60°57.226'N 138°02.466'W, P. Caswell 03-037, 12 
June 2003 (DAO); dry organic soil, middle of road, 
abandoned road from Dalton Post to Wade Lakes, 
60°06.192'N 137°07.847'W, P. Caswell 24, 6 June 
2002 (DAO) (determined by G. A. Mulligan). 

The first specimen cited above is from a site intermediate 
between two areas in the Park mapped by Cody (1996). The 
second specimen is the most southerly yet found in the Park 


about 30 kilometers south of a site reported in Cody et al. 
(2003). 


Draba cinerea J. E. Adams, Gray-leaved Whitlow- 
grass — YUKON: Kluane National Park, moist organic 
soil on hill above an alpine marsh, nunatak at Logan 
Warden Cabin, 60°47.598'N 140°45.588'W, P. Caswell 
et al. 03-32S8A, 12 July 2003 (DAO) (determined by 
G. A. Mulligan). 

The specimen cited above is the most southwesterly yet 


found in the Territory. It is from a site about 50 kilometers 
south of a site at about latitude 61°10'N (Cody 1996). 


Draba crassifolia Grah., Rocky Mountain Whitlow- 
grass — YUKON: Kluane National Park, organic soil 
among large rocks, old lake bed, nunatak at Logan 
Warden Cabin, 60°47.500'N 140°44.991'W, P. Caswell, 
R. Maraj & L. Fries 03-291, 12 July 2003 (DAO) 
(determined by G. A. Mulligan). 

The specimen cited above is an extension of the known 
range in the Park of about 80 kilometers west of two sites 
mapped by Cody (1996) and is the westernmost yet known 
in that area. 


Draba lonchocarpa Rydb. var. vestita O. E. Schulz, 
Alaska Whitlow-grass — YUKON: Kluane National 
Park, rocks and gravel, Cache Lake, 61°12.454'N 
139°03.757'W, P. Caswell 60B, 12 June 2002 (DAO) 
(determined by G. A. Mulligan); Kluane National Park, 
glacial till with thin organic cover, nunatak at Logan 
Warden Cabin, 60°47.628'N 140°45.148'W, P. Caswell 
et al. 03-351, 12 July 2003 (DAO) (determined by G. 
A. Mulligan). 

Cody et al. (2001) reported the first record of this rare 
variety in the Territory from the extreme southeast. The 
specimens cited above extend the known range in the Terri- 
tory about 1000 kilometers to the west to near the Alaska 
border. 


Draba nivalis Liljebl., Snow Draba — YUKON: Kluane 
National Park, moist organic soil on hill above an 
alpine marsh, nunatak at Logan Warden Cabin, 
60°47.598'N 140°45.588'W, P. Caswell et al. 03- 
328A-B, 12 July 2003 (DAO) (determined by G. A. 
Mulligan). 

The specimen cited above is the most southwesterly yet 


found in the Territory. It is from a site about 100 kilometers 
west of a site at about 138°30'W (Cody 1996). 


Draba oligosperma Hook., Few-seeded Draba, Few- 
seeded Whitlow-grass — YUKON: scree slope, dry rocky 
soil with little humus, abandoned road between Dal- 
ton Post and Wade Lakes, 60°05.992'N 137°08.531'W, 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


P. Caswell 02-25, 6 June 2002 (DAO) (determined by 
G. A. Mulligan). 

The specimen cited above which is the southernmost yet 
found in Kluane National Park is an extension of the known 
range in the Territory of about 75 kilometers southeast of a 
site southwest of Haines Junction. 


Draba scotteri G. A. Mulligan — YUKON: Kluane 
National Park, scree with organic soil, alpine tundra, 
nunatak at Logan Warden Camp, 60°46.958'N 140° 
44.864'W, P. Caswell et al. 284, 11 July 2003 (DAO); 
moist organic soil alpine tundra on hillside, nunatak 
at Logan Warden Cabin, 60°47.545'N 140°45.334'W, 
P. Caswell et al. 03-176, 10 July 2003 (DAO) (deter- 
mined by G. A. Mulligan). 

This is a rare species in the Territory (Douglas et al. 
1981). The specimens cited above are from sites about 120 
kilometers west of sites mapped by Cody (1996) in Kluane 
National Park. 


Draba_ stenoloba Ledeb., Alaska Draba — YUKON: 
Kluane National Park, organic soil, right of way of 
abandoned pipeline, south of Auriol Trail, 60°42.932'N 
137°25.565'W, P. Caswell 03-70, 21 June 2003 (DAO) 
(determined by G. A. Mulligan). 

The specimen cited above is from a site intermediate 
between sites mapped by Cody (1996) on the Haines and 
Alaska highways. 


Draba stenopetala Trautv., Starflowered Whitlow-grass 
— YUKON: small quantity of inorganic soil between 
two rocks, barren mountain summit, mountain east of 
Haines Hwy. just north of British Columbia border, 
60°03.115'N 136°50.004'W, L. Struever s.n., 1 July 
2003 (DAO) (determined by G. A. Mulligan). 

Douglas et al. (1981) knew this rare species in the Terri- 
tory from only three sites in Kluane National Park. Cody 
(1996) mapped two additional sites east of the Dempster Hwy. 
and Cody et al. (2001) added two additional sites in the 
south at longitudes 135°29'W and 134°41'W. The specimen 
cited above is from a site about 30 kilometers east of the 
nearest site in southern Kluane National Park. 


DROSERACEAE 
Drosera anglica Huds., Great Sundew — YUKON: 
growing at edge of small thermokarst pond, Alaska 
Hwy. 6 km NW of Dry Creek #2, 62°13.455'N 140° 
41.544'W, B.A. Bennett, R. Elven & H. Solstad 03-046, 
8 Aug. 2003 (B. A. Bennett Herbarium, photo DAO). 
This species was considered rare in the Territory by Dou- 
glas et al. (1981). The specimen cited above is from a site 
about 250 kilometers southwest of a site in the vicinity of 
Mayo (Cody 1996). 


SAXIFRAGACEAE 

Leptarrhena_ pyrolifolia (D.Don) R.Br. ex Ser., 
Leather-leaved Saxifrage — YUKON: Kluane National 
Park, Fisher Glacier near Alsek River, lower south- 
facing slope north of glacier, 60°08'31"N 138°13'13"W, 
B. A. Bennett 03-978, 6 July 2003 (Yukon Government 
Herbarium, photo DAO). 


2005 


Cody (1996) knew this species in the Park from only two 
sites about halfway between this site and Haines Junction. 
The nearest other site was about 300 kilometers to the 
northeast adjacent to the South Canol Road. 


Parnassia fimbriata Koenig, Fringed Grass-of-Par- 
nassus — YUKON: Kluane National Park, Fisher Glaci- 
er near Alsek River, NE of base camp, common in 
open herbaceous and graminoid meadows of south- 
facing lower slopes north of camp, 60°08'31"N 138° 
13'13"W, B. A. Bennett 03-1066, 8 July 2003 (DAO). 

The specimen cited above is the southernmost yet found 
in the Park. It is about 50 kilometers southeast and south- 
west of the only two other sites mapped by Cody (1996) in 
the Park. 


Saxifraga nelsoniana D.Don ssp. carlottae (Calder & 
Savile) Hultén (S. punctata L. ssp. carlottae Calder 
& Savile) — YUKON: in wet sand and gravel by stream- 
let on south side of large pond at Long Warden 
Cabin, nunatak, 60°47.628'N 140°45.148'W, P. Caswell 
et al. 03-368, 13 July 2003 (DAO). 

The specimen cited above fits this subspecies which is 
new to the Yukon Territory (Cody 1996) on the basis of the 
scarcely cordate leaves with less than 12 lobes. It is, however, 
in flower and lacks the long, narrow, deeply cleft capsules. 
The following information is provided from Calder and 
Savile (1960): “Subspecies carlottae is abundant on moist 
alpine or subalpine slopes or clefts in the Queen Charlotte 
Islands; but it has also spread to the mainland where it inter- 
grades freely with ssp. pacifica and ssp. porsildiana. ... 
Attention must be drawn to the plants in the Prince William 
Sound region of southern Alaska, cited as intermediate 
between ssp. carlottae and ssp. pacifica. These plants are 
actually closer in morphology to carlottae than pacifica, 
despite the occurrence of pure pacifica further east.” 


Saxifraga nelsoniana D.Don ssp. pacifica (Hultén) 
Hultén (Saxifraga punctata L. ssp. pacifica Hultén) — 
YUKON: Kluane National Park, Fisher Glacier near 
Alsek River, W-facing slope above valley glacier, 
60°08'25"N 138°13'20"W, B. A. Bennett 03-1001, 7 
July 2003 (DAO). 

The specimen cited above is the most southwesterly yet 
found in the Park. It is from a site about 75 kilometers west 
of a site adjacent to the Haines Highway. 


Saxifraga rufopilosa (Hultén) A. E. Porsild — YUKON: 
Kluane National Park, Fisher Glacier near Alsek River, 
W-facing slope above valley glacier, 60°08'20"N 
138°13'20"W, B. A. Bennett 03-996, 7 July 2003 (DAO). 
The specimen cited above is the southernmost yet found 
in the Park. It is from a site about 60 kilometers south of a 
site mapped by Cody (1996) southwest of Haines Junction. 


ROSACEAE 

Dryas integrifolia Vahl ssp. crenulata (Juz.) J. Kozhevn 
(D. crenulata Juz.) — YUKON: Kluane National Park, 
Fisher Glacier near Alsek River, base camp, common 
on south and west-facing slopes and occasionally on 
upper slopes and in valley, 60°08'31"N 138°13'13"W, 
B. A. Bennett 03-978, 6 July 2003 (B. A. Bennett 
Herbarium, photo DAO). 


Copy, BENNETT, AND CASWELL: VASCULAR PLANTS IN THE YUKON VII 


43] 


The specimen cited above is the southernmost yet found 
in the Park. It is from a site about 100 kilometers south of a 
site mapped by Cody (1996) southeast of Kluane Lake. 


Dryas octopetala L. ssp. hookeriana (Juz.) Hultén 
(D. hookeriana Juz.) — YUKON: Kluane National Park, 
Fisher Glacier near Alsek base camp, common on 
south- and west-facing slopes, 60°08'31"N 138°13'13"W, 
B. A. Bennett 03-1085, 6 July 2003 (B. A. Bennett 
Herbarium, photo DAO). 

The specimen cited above is the southernmost yet known 
in the Park. It is about 30 kilometers south of a site south- 
west of Haines Junction mapped by Cody (1996). 


Luetkea pectinata (Pursh) Kuntze, Partridge-foot — 
YUKON: Kluane National Park, Fisher Glacier near 
Alsek River, west-facing slope above valley glacier 
leading into Fisher Glacier, 60°08'16"N 138°13'28"W, 
B. A. Bennett 03-988, 7 July 2003 (Yukon Government 
Herbarium, photo DAO). 

This species was known to Cody (1996) from only two 
sites in the southeast of the Park. The specimen cited above 
is an extension of the known range in the Park of about 25 
kilometers west of the southernmost site. 


Potentilla bipinnatifida Doug]. ex Hook., Bipinnate 
Cinquefoil — YUKON: Kluane National Park, packed 
earth and gravel with organic component, centre of road, 
Alsek Trail, P, Caswell 02-437, 12 July 2002 (DAO). 

Cody (1996) stated that this species was apparently rare 
in northern and southwestern parts of the Territory but it 
was not included in The Rare Vascular Plants of the Yukon 
Territory (Douglas et al. 1981). The specimen cited above is 
from a site intermittent between two sites mapped by Cody 
in the Park. 


Rosa woodsii Lindl., Western Rose — YUKON: Pelly 
Crossing, growing on upper south-facing 45°bluff 
slope, 62°49'S7"N 136°34'45"W, B. A. Bennett 03- 
095, 1 Sept. 2003 (DAO). 

The specimen cited above is an extension of about 90 
kilometers north of Carmacks. 


Sorbus sitchensis Roemer — YUKON: Kluane National 
Park, Fisher Glacier near Alsek River, NE of base 
camp, only a single plant seen in open herbaceous and 
graminoid meadows of south-facing lower mid-slope 
growing 1.5 meters tall, 60°08'31"N 138°13'13"W, 
B. A. Bennett 03-1071, 8 July 2003 (DAO). 

This species which was considered rare in the Territory 
by Douglas et al. (1981) was known to Cody (1996) from only 
two nearby sites in the extreme south of Kluane National 
Park. The specimen cited above is an extension of the known 
range in the Territory of about 25 kilometers to the west. 


LEGUMINOSAE (FABACEAE) 
Astragalus alpinus L., Alpine Milk-vetch — YUKON: 
Kluane National Park, nunatak between Lowell and 
Dusty glaciers, near Ulu Mountain, 60°21'32"N 138° 
34'49"W, B. A. Bennett 03-941, 6 July 2003 (DAO). 
The specimen cited above is from the most southwesterly 
site in the Park. This site is about 25 kilometers west of a 
site mapped by Cody (1996). 


432 


Lupinus nootkatensis Donn ex Sims, Nootka Lupine 
— YUKON: Kluane National Park, Fisher Glacier near 
Alsek River, slope above valley glacier leading into 
the glacier, 60°08'16"N 138°28'W, B. A. Bennett 03- 
1006, 7 July 2003 (Yukon Government Herbarium, 
photo DAO). 

The specimen cited above is an extension of the known 
range in the Park of about 25 kilometers southwest of this rare 
species in the Territory (Douglas et al. 1981) which was 
known to Cody (1996) from only two sites in the extreme 
south and an additional site adjacent to the Haines Highway 
(Cody et al. 2004). 


Oxytropis arctica R.Br. var. arctica, Arctic Oxytrope 
— YUKON: Vuntut National Park, ca. 13 miles W of 
Sam Lake, 68.35°N 139.1667°W, S. L. Welsh 10340, 
9 July 1970 (ALA). 

Cody (1996) described and keyed this species, which is 
known from arctic Canada and northern Alaska, and sug- 
gested that it should be looked for in the mountains of 
northern Yukon Territory. It should be added to the list of 
rare plants in the Territory (Douglas et al. 1981). 


Oxytropis arctica R.Br. var. murrayi (Yurtzev) S. 
Welsh (QO. sordida [Willd.] Pers. ssp. murrayi B. A. 
Yurtsev), Murray’s Oxytrope — YUKON: Kluane Nation- 
al Park, Observation Mtn. and vicinity at terminus of 
Kaskawulsh Glacier, 60.8167°N 138.7334°W, D. F: and 
B. Murray 522, 10 July 1966 (ALA); Kluane National 
Park, Fisher Glacier near Alsek River, west-facing 
15° slope, above valley glacier in Dryas integrifolia 
heath tundra with Hedysarum boreale, Senecio ogor- 
torukensis, some limestone in the thin soil, just near- 
ing ridge, 60°08'12"N 138°13'14"W, B. A. Bennett 
03-981, 7 July 2003 (DAO). 

This plant was fairly common on this site but became 
less common quickly as you follow the ridge to the summit. 
This is a new variety first reported in Cody et al. (2004) in 
flora of the Yukon Territory, which is endemic to Kluane 
National Park. It should be added to the list of rare plants 
(Douglas et al. 1981). Oxytropis arctica var. murrayi can be 
separated from var. arctica as follows: 

A. Leaves seldom with some leaflets 

fasciculate; leaflets often over 8 mm 

long, at least some, plants of arctic 

Dea Keay) hee SP Re Beek yh at les var. arctica 
A. Leaves typically with fasciculate 

leaflets; leaflets mainly less than 

8 mm long; plants of SW Yukon ....... var. murrayi 

Oxytropis arctica var. murrayi can be described as follows: 
Plants mainly 4-16 cm tall. Leaves 3-12 cm long; leaflets 
17-numerous, opposite, scattered or more typically fascicu- 
late, mainly 3-8 mm long, but in some up to 14 mm. Scapes 
4-12 (14) cm; racemes 2-7 flowered, subcapitate or slightly 
elongate; calyx tube shaggy villous with light and dark hairs, 
4-5 mm long. Pods 17-21 mm, spreading. Type: Yukon, St. 
Elias Mts., Observation Mt. and vicinity, at terminus of 
Kaskawulsh Glacier, D. F’ & B. M. Murray 522, 10 July 1966. 
Flowering in summer. Ridge crests, talus slopes, and meadows 
at 725-1830 m. 


Oxytropis campestris (L.) DC. ssp. jordalii (A. E. 


Porsild) Hultén, Jordal’s Oxytrope — YUKON: Kluane 
National Park, Fisher Glacier near Alsek River, NE 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


of base camp, drier open south-facing mid slope, 
60°08'31"N 138°13'13"W, B. A. Bennett 03-1070, 8 
July 2003 (DAO); west-facing slope 40°-50°, above 
valley glacier leading into Fisher Glacier, 60°08'16"N 
138°13'28"W, B. A. Bennett 03-992, 7 July 2003 (B. 
A. Bennett Herbarium, photo DAO). 

The specimens cited above are the first reported from 
Kluane National Park and are an extension of the known 
range in the Territory of about 200 kilometers west of sites 
in the vicinity of Carcross (Cody 1996). 


Oxytropis nigrescens (Pall.) Fisch. ssp. nigrescens, 
Blackish Locoweed — YUKON: Kluane National Park, 
Fisher Glacier near Alsek River, west-facing slope 
40°-50° above valley glacier leading into Fisher Glacier, 
60°08'16"N 138°13'28"W, B. A. Bennett 03-991, 7 July 
2003 (DAO). 

The specimen cited above is an extension of the known 
range in the Park of about 75 kilometers south of a site 
mapped by Cody (1996) west of Haines Junction. 


Vicia americana Muhl., American Vetch — YUKON: 
behind cabins, north end of Dawson City by White- 
horse Cabins, 64°04'N 139°26'W, G. Brunner 585, 
12 Sept. 2003 (DAO). 

The collection cited above is from a site about 175 kilo- 
meters west-northwest of Mayo and was undoubtedly intro- 
duced there. 


GERANIACEAE 

Geranium erianthum DC., Northern Geranium — 
YUKON: Kluane National Park, organic soil in meadow, 
Plug Creek, 60°03.593'N 138°12.055'W, P. Caswell 
et al. 03-675, 29 July 2003 (DAO). 

This is a rare species in the Territory (Douglas et al. 1981) 
where it is mainly found in Kluane National Park. The spec- 
imen cited above is the westernmost site adjacent to the 
British Columbia border. 


VIOLACEAE 
Viola adunca J. E. Smith, Hook-spur Violet — YUKON: 
Kluane National Park, growing in shady areas under 
willows by small creek on west slope above valley gla- 
cier leading into Fisher Glacier near Alsek River, 
60°08'10"N 138°13'15"W, B. A. Bennett 03-985, 7 
July 2003 (DAO). 

The specimen cited above is an extension of the known 
range in the Park of about 35 kilometers northwest of a site 
mapped by Cody (1996) near the British Columbia border. 


ONAGRACEAE 
Epilobium hornemannii Reichenb. ssp. hornemannii, 
Hornemann’s Willowherb — YUKON: Kluane National 
Park, dry wind-packed fine scree, nunatak north of 
Lowell Glacier, 60°18.297'N 138°34.503'W, P. Caswell 
03-604, 29 July 2003 (DAO). 

The specimen cited above is an extension of the known 
range into southern Kluane National Park of about 100 kilo- 


meters southwest of a site east of Haines Junction (Cody 
1996). 


HALORAGACEAE 
Myriophyllum verticillatum L., Verticillate Water-mil- 
foil — YUKON: muddy substrate, in two feet of water, 


2005 


growing in dense clumps, Poison Lake, 60°14.686'N 
136°57.421'W, P. Caswell 03-493, 24 July 2003 (DAO). 

This is a rare species in the Territory (Douglas et al. 1981). 
The specimen cited above is an extension of the known range 
of about 80 kilometers southeast of a site northwest of Haines 
Junction (Cody 1996). 


APIACEAE 

Angelica lucida L., Seacoast Angelica — YUKON: Klu- 
ane National Park, moist, organic soil, Plug Creek, 
60°03.593'N 138°12.055'W, P. Caswell et al. 03-666, 
29 July 2003 (DAO). 

This is a rare species in the Yukon Territory (Douglas et 
al. 1981). The specimen cited above is the westernmost yet 
known in the Territory and is an extension of about 75 kilo- 
meters west of the Lower Alsek River in the extreme south- 
east of the Park (Douglas and Ruyle-Douglas 1978). 


ERICACEAE 
Harrimanella stellariana (Pallas) Coville (Cassiope 
stellariana (Pallas) DC.), Alaskan Mountain-heather 
— YUKON: Kluane National Park, abundant sometimes 
forming nearly pure stands covering areas in the lower 
valley and the lower north-facing slopes, Fisher Glac- 
ier near Alsek River, base camp, 60°08'31"N 138° 
13'13"W, B. A. Bennett 03-953, 6 July 2003 (DAO). 
This species was reported as rare in the Territory by 
Douglas et al. (1981) on the basis of only three known sites, 
two of which were in Kluane National Park. The specimen 
cited above is an extension of about 25 kilometers to the 
west in the Park. 


Phyllodoce X intermedia (Hook.) Rydb. — YUKON: 
Kluane National Park, Fisher Glacier near Alsek River, 
southwest-facing slope 30° above Fisher Glacier, 
60°08'06"N 138°12'58"W, B. A. Bennett 03-1035, 7 
July 2003 (B. A. Bennett Herbarium, photo DAO). 

Cody et al. (2001) reported the first known occurrence of 
this hybrid between Phyllodoce empetriformis and P. glan- 
dulifera from a wooded site near Kathleen Lake at about Mile 
140 Haines Road, about 75 kilometers northwest of the sec- 
ond site in Kluane National Park. 


PRIMULACEAE 
Douglasia ochotensis (Willd.) Hultén, Arctic-Mon- 
tane Dwarf-Primrose — YUKON: Dempster Highway 
kilometer 440, 4 km southwest of Rock River Camp- 
ground with Salix phlebophylla in platey loose shale 
slope, 66°52'37"N 136°19'58"W, B. A. Bennett 03- 
078, 13 Aug. 2003 (DAO). 

The specimen cited above is the easternmost yet found in 
the Territory. It is from a site about 175 kilometers south- 
east of the nearest site mapped by Cody (1996). 


Primula nutans Georgi (P. sibirica Jacq.), Siberian 
Primrose — YUKON: damp organic soil, uneven tundra, 
Alaska Hwy. at squirrel research area, 60°57.156'N 
138°02.086'W, P. Caswell 03-034, 12 June 2003 (DAO). 

This Amphi-Beringian species is found in North America 
in extreme western Alaska and then disjunct to eastern Alas- 
ka and southwestern Yukon Territory (Hultén 1968). The spec- 
imen cited above is a slight extension to the northwest from 
sites in the Haines Junction region mapped by Cody (1996). 


Copy, BENNETT, AND CASWELL: VASCULAR PLANTS IN THE YUKON VII 


433 


GENTIANACEAE 
Swertia perennis L., Alpine Bog Swertia — YUKON: wet 
organic soil, open Picea glauca forest, second gully 
east of Haines Highway proceeding north from the 
British Columbia border, 60°00.403'N 136°51.061'W, 
P. Caswell 03-546, 2 Aug. 2003 (DAO). 

Cody (1996) suggested that this species was to be looked 
for in southwestern Yukon Territory. It has now been found 
and should be added to the list of rare plants in the Territory. 


MENYANTHACEAE 
Menyanthes trifoliata L., Buckbean, Bogbean — YUKON: 
pond west of Alaska Hwy., north of Donjek bridge, 
61°41'11.8"N 139°47'01.6"W, D. Normadeau s.n., 26 
July 2003 (DAO); emergent in highway pond, 4 km 
SE of Pickhandle Lake, 61°54'20"N 140°13'00"W, B. 
A. Bennett 97-27, 6 June 1996 (DAO); “Eagle Lake’, 
61°23'30"N 139°41'30"W, R. D. Wickstrom s.n., 20 
Aug. 1974 (DAO). 

The three sites cited above extend the known range in the 


Territory about 115 kilometers southeast of a site adjacent 
to the Alaska Hwy. and Alaska border. 


APOCYNACEAE 
Apocynum androsaemifolium L., Spreading Dogbane 
— YUKON: in open dry Pinus contorta/lichen woods, 
Watson Lake area, 60°11'N 129°O5'W, R. Rosie 1893, 
6 July 1995 (DAO); open Pine/Lichen bluff above lake, 
Blind Lake, 60°04'43.3"N_ 128°15'3.7"W, J. Stani- 
forth 2, 26-28 June 2003 (DAO). 

Cody (1996) knew this species as occasional north to the 
vicinity of Dawson City. The specimens cited above are from 
sites west of the easternmost mapped by Cody. 


POLEMONIACEAE 
Polemonium pulcherrimum Hook., Showy Jacob’s 
Ladder — YUKON: Kluane National Park, nunatak 
between Lowell and Dusty glaciers near Ulu Moun- 
tain, on 35-40° south-facing slope, 60°21'32"N 138° 
34'49"W, B. A. Bennett 03-923, 6 July 2003 (DAO). 
The specimen cited above is from a site intermediate 
between two sites in the extreme south of the Park about 
125 kilometers apart. 


VERBENACEAE 

Verbena hastata L. — A specimen which was collected by 
Phil Caswell at Haines Junction on 13 August 2000 was 
identified to this species by B. A. Bennett and confirmed by 
W. J. Cody. It was reported as new to the Yukon Territory by 
Cody et al. (2003). Unfortunately this specimen was mis- 
identified and has been correctly revised to Veronica longi- 
folia L., by Stuart G. Hay of the Herbier Marie-Victorin 
(MT), Université de Montréal, a species which was first col- 
lected along the Alaska Hwy. by Cody and Ginns in July 
1980. Verbena hastata should be deleted from the flora of 
the Yukon Territory. 


SCROPHULARIACEAE 

Castilleja miniata Doug}. ex Hook., Scarlet Paintbrush 
— YUKON: Kluane National Park, moist, organic soil, 
meadow on north shore of first lake from St. Elias 
Lake, Secret Lakes, 60°18.979'N 137°08.208'W, P. 
Caswell & L. Freese 03-524, 26 July 2003 (DAO). 


434 


This species was considered rare in the Territory by Dou- 
glas et al. (1981) who knew it only east of Atlin Lake and 
the Larson Creek hot springs. Catherine Kennedy collected 
it at the Coal River Springs area in 1983 (Cody 1994) and 
Mark Egger collected it adjacent to the Top of the World 
Hwy. west of Dawson City in 1991 (Egger 1992). The spec- 
imen cited above is a new record for Kluane National Park, 
about 200 kilometers west of the site near Atlin Lake. 


Castilleja parviflora Bong., Small-flowered Paint- 
brush — YUKON: Kluane National Park, scree with a 
small organic component, nunatak (abandoned weath- 
er station) north of Lowell Glacier, 60°18.297'N 
138°34.503'W, P. Caswell et al. 03-603, 29 July 2003 
(DAO). 

This rare plant in the Territory (Douglas et al. 1981) is only 
known in the Territory in the southeast of Kluane National 
Park. The specimen cited above is from a site about 75 kilo- 


meters northwest by west of the southernmost mapped by 
Cody (1996). 


Castilleja unalaschcensis (Willd.) Hultén, Unalaska 
Paintbrush — YUKON: Kluane National Park, Fisher 
Glacier near Alsek River, southwest-facing 30° slope 
above Fisher Glacier, 60°08'06"N 138°12'58"W, B. 
A. Bennett 03-1021, 7 July 2003 (DAO). 

The specimen cited above is an extension of the known 
range in the southern part of the Park of about 30 kilometers 
west of a site mapped by Cody (1996). 


Veronica americana Schwein., American Brooklime 
— YUKON: Haines Hwy., interface of dried mud and 
water, beaver pond on east side of highway about 10 km 
south of Kathleen Lake cutoff, 60°30.838'N 137° 
04.970'W, P. Caswell 02-788, 11 Aug. 2002 (DAO). 

Cody (1996) reported this species as occasional in the 
Territory north to about latitude 63°N. The specimen cited 
above is from a site about 80 kilometers southeast of the 
westernmost location in the Territory adjacent to the Alaska 
Hwy. 


OROBANCHACEAE 

Orobanche fasciculata Nutt., Clustered Broomrape — 
YUKON: mid-slope, Conglomerate Mountain, 61.6252°N 
135.8755°W, B. A. Bennett & S. Thompson 03-011, 2 
May 2003 (DAO); old esker complex with silty soil, 
gravel and boulders at crest of south-facing slope, 
Haunka Creek, 60°14'05.4"N_ 133°53'29.3"W, B. A. 
Bennett, R. & P. Mulder 04-0052, 22 May 2004 
(DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory. The first specimen cited above is from a site inter- 
mediate between Carmacks and a site just north of White- 
horse (Cody 1996). The second specimen cited above is an 
extension of about 30 kilometers to the southeast of a site 
mapped by Cody (1996) adjacent to the Alaska Hwy. and is 
the southernmost yet known in the Territory. 


LENTIBULARIACEAE 

Utricularia minor L., Lesser Bladderwort — YUKON: 

Kluane National Park, in five inches water, muddy 

substrate, pond at Slims East Trailhead, 60°59.408'N 

138°29.483'W, P. Caswell 03-541, 28 July 2003 (DAO). 
The specimen cited above is an extension of the known 

range of this uncommon species in the Territory of about 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


300 kilometers west of a site near the south end of the Canol 
Road (Cody 1996). It is new to Kluane National Park. 


PLANTAGINACEAE 
Plantago eriopoda Torr., Saline Plantain — YUKON: 
dry gravel, edge of parking space, Burwash Landing 
Airport, 61°22.183'N 139°01.842'W, P. Caswell 02- 
319, 6 July 2002 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 75 kilometers northwest of a 


site adjacent to the Alaska Hwy. northwest of Haines Junc- 
tion (Cody 1996). 


RUBIACEAE 
Galium triflorum Michx., Sweet-scented Bedstraw — 
YUKON: organic soil at edge of thicket on west-facing 
slope, east of Haines Hwy. at Km 152, 60°02.970'N 
136°53.001'W, P. Caswell 03-746, 21 Aug. 2003 (DAO). 
This is a rare species in the Territory (Douglas et al. 1981). 
The specimen cited above is an extension of the known 
range of about 140 kilometers southeast of a site near the 
south end of Kluane Lake. 


CAPRIFOLIACEAE 
Lonicera involucrata (Richards.) Banks ex Spreng., 
Black Twinberry — YUKON: Haines Junction, L. Fournier 
s.n., July 1958 (DAO). 

The specimen cited above was likely collected from cul- 
tivated plants. In the Yukon, it is known only from persist- 
ing plantings. 


VALERIANACEAE 
Valeriana sitchensis Bong., Sitka Valerian — YUKON: 
Kluane National Park, moist organic soil, Plug Creek, 
60°03.593'N 138°12.055'W, P. Caswell 03-647, 29 
July 2003 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 25 kilometers to the south- 
west of a site mapped by Cody (1996). 


CAMPANULACEAE 

Campanula rotundifolia L., Harebell — YUKON: Klu- 
ane National Park, open soil with Festuca altaica, 
Arcostaphylos uva-ursi, Juniperus communis, Shep- 
herdia canadensis, Populus balsamifera, and Potentilla 
fruticosa, Fisher Glacier near Alsek River, 60°07'54"N 
138°12'23"W, B. A. Bennett 03-1052, 8 July 2003 (B. 
A. Bennett Herbarium, photo DAO). 

This species was considered rare in the Territory by Dou- 
glas et al. (1981). The specimen cited above is from the third 
site in Kluane National Park and is an extension of about 30 
kilometers west of a site mapped by Cody (1996). 


ASTERACEAE 
Antennaria pulcherrima (Hook.) Greene, Showy 
Pussytoes — YUKON: Watson River, near mouth by 
Carcross Dunes, 60°11'05"N 134°44'16"W, B. A. 
Bennett & R. Mulder 03-1308, 17 Aug. 2003 (DAO). 
The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 125 kilometers 
west of a site in the vicinity of Teslin. 


Arnica diversifolia Greene, Diverse Arnica — YUKON: 
Kluane National Park, scree with organic component, 
mountain slope, old weather station on nunatak north 


2005 


of Lowell Glacier, 60°18.297'N 138°34.503'W, P. 
Caswell et al. 03-597, 29 July 2003 (DAO). 

This is a rare species in the Territory (Douglas et al. 
1981) where it was known from a site near the south end of 
Kluane Lake in Kluane National Park. The specimen cited 
above is a southward extension of the known range in the 
Park of about 75 kilometers. 


Arnica latifolia Bong., Mountain Arnica — YUKON: 
Kluane National Park, Fisher Glacier near Alsek River, 
45° southwest-facing slope above Fisher Glacier, silty 
sandy soil, 60°08'03"N 138°12'54"W, B. A. Bennett 
03-1033, 7 July 2003 (DAO). 

The specimen cited above is the fourth yet known in the 


Park and the most westerly yet known in the Territory (Cody 
1996). 


Artemisia tilesii Ledeb. s.l., Aleutian Mugwort — 
YUKON: Kluane National Park, occasional on xeric 
rock outcrops in shallow soil on edge of extremely steep 
cliffs, also seen at base of mountain, Fisher Glacier 
near Alsek River, 60°08'06"N 138°12'42"W, B. A. Ben- 
nett 03-1022, 7 July 2003 (DAO). 

The specimen cited above is an extension of the known 
range in the Park of about 30 kilometers to the west from a 
site in the extreme south. This specimen is a very distinct form 
with rounded leaflets and dark bracts and may represent a 
new variety. 


Cirsium foliosum (Hook.) DC., Leafy Thistle, Elk 
Thistle — YUKON: Beaver Pond on McIntyre Creek 
across from Kopper King, Whitehorse, 60°44.73'N 
135°07.36'W, G. Delaet, B. A. Bennett & R. Mulder 
03-1370, 29 Sept. 2003, (B. A. Bennett Herbarium, 
photo DAO). 

The specimen cited above is from a site intermediate 


between sites adjacent to the South Canol Road and Haines 
Hwy. mapped by Cody (1996). 


Erigeron grandiflorus Hook. ssp. arcticus A. E. Por- 
sild, Large-flowered Daisy — YUKON: Kluane National 
Park, organic soil on south-facing slope of alpine 
meadow, Wade Mountain, 61°18.373'N 139°31.273'W, 
P. Caswell 02-374, 7 July 2002 (DAO). 

This species was not included in The Rare Plants of the 
Yukon (Douglas et al. 1981) because of its widespread range. 
The specimen cited above is a short distance northwest of 
the westernmost site mapped by Cody (1996) in the Kluane 
National Park. 


Erigeron humilis Graham, Arctic-alpine Daisy — YUKON: 
Kluane National Park, south-facing alpine slope, 
nunatak (abandoned weather station) north of Lowell 
Glacier, 60°18.297'N 138°34.503'W, P. Caswell 03- 
591, 29 July 2003 (DAO). 

The specimen cited above is the most southwesterly yet 
known in Kluane National Park (Cody 1996). The nearest 
known site is about 25 kilometers to the northeast. 


Erigeron peregrinus (Banks ex Pursh) Greene ssp. 
peregrinus, Subalpine Daisy — YUKON: Kluane Nation- 
al Park, moist organic soil, meadow, Plug Creek, 60° 
03.593'N 138°12.055'W, P. Caswell et al. 03-651, 29 
July 2003 (DAO). 

This subspecies was considered rare in the Territory by 
Douglas et al. (1981). The specimen cited above is an ex- 


Copy, BENNETT, AND CASWELL: VASCULAR PLANTS IN THE YUKON VII 


435 


tension of the known range in the Territory of about 30 kilo- 
meters west of a site near the British Columbia border. 


Erigeron pumilus Nutt., Shaggy Fleabane — YUKON: 
Kluane National Park, Donjek Valley, 61°00'N 139° 
22.573'W, R. Maraj s.n., 15 July 2002 (DAO). 

Douglas et al. (1981) reported this species as rare in the 
Yukon Territory on the basis of a specimen collected in the 
vicinity of Kluane Lake. The specimen cited above is only 
the second known in the Territory. It was collected about 65 
kilometers to the southwest and is a new record for Kluane 
National Park. 


Erigeron uniflorus L. ssp. eriocephalus (Vahl ex 
Hornem.) Cronq., Northern Daisy — YUKON: Kluane 
National Park, alpine scree slope, nunatak at Logan 
Warden Cabin, 60°46.889'N 140°45.108'W, P. Caswell 
et al. 03-264, 11 July 2003 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory. The specimen cited above which is about 50 kilo- 
meters southwest of the westernmost mapped by Cody (1996) 
in Kluane National Park is now the westernmost yet known 
in the Park. 


Erigeron yukonensis Rydb., Yukon Fleabane — YUKON: 
organic soil, flat burned area of dead Picea glauca, 
Copper Joe Creek, 61°18.641'N 138°56.324'W, P. 
Caswell 02-156, 24 June 2002 (DAO); barren ridge, 
50% vegetative coverage, east-facing slope 2°, inorgan- 
ic soil with Dryas intergrifolia ssp. intergrifolia and 
Saxifraga tricuspidata, Thunder Egg Mountain, col- 
lected by Z. Mattson 60°47.777'N 137°46.993'W, P. 
Caswell 04-058 (Kluane Park Herbarium, photo DAO). 

This species is an endemic of the Yukon Territory and 
adjacent District of Mackenzie (Cody 1996). The first spec- 
imen cited above is an extension of the known range in the 
Territory of about 100 kilometers northwest of a site in Klu- 
ane National Park southwest of Haines Junction. 


Hieracium gracile Hook., Slender Hawkweed — 
YUKON: Kluane National Park, Fisher Glacier near 
Alsek River, west-facing slope above valley glacier 
leading into Fisher Glacier, 60°08'16"N 138°13'28"W, 
B. A. Bennett 03-987, 7 July 2003 (DAO). 

The specimen cited above is an extension of the known 
range about 35 kilometers west of the southernmost previ- 
ously known to Cody (1996) in the Park. 


Symphyotrichum ciliatum (Lindl.) A. & D. Love (Aster 
brachyactis Blake) Rayless Aster, Lindley’s American 
Aster — YUKON: growing in silty saline mud along east 
shore and amongst Puccinellia, Triglochin palustris, 
Triglochin maritima and Lomatagonium rotatum, Fox 
Creek saline flat 600 m south of Fox Creek, Km 227 
on east side of North Klondike Hwy., B. A. Bennett & 
R. Mulder 03-1341, 13 Sept. 2003 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory on the basis of a single site at an alkaline meadow 
adjacent to the Takhini River west of Whitehorse. The spec- 
imen cited above from north of Whitehorse is from only the 
second known site in the Territory. 


Symphyotrichum yukonense (Cronquist) Nelson (Aster 
yukonensis Cronquist), Yukon Aster, Yukon-American 
Aster — YUKON: silty broad alkaline river flat, north- 
west side of Donjek River, 61°40.8'N 139°45.58'W, 


436 


B. A. Bennett, R. Elven & H. Solstad 03-053, 8 Aug. 
2003 (DAO). 

This species was considered rare in the Territory by Dou- 
glas et al. (1981). The specimen cited above is an extension 
of the known range in the Territory of about 100 kilometers 
northwest of sites in the vicinity of the south end of Kluane 
Lake. 


Taraxacum lyratum (Ledeb.) DC., Rock Dandelion — 
YUKON: Kluane National Park, Fisher Glacier near 
Alsek base camp, occasional in valley bottom in 
heath tundra, 60°08'31"N 138°13'13"W, B. A. Ben- 
nett 03-956, 6 July 2003 (DAO). 

The specimen cited above is from the southernmost site 
yet found in the Park. It is from a site about 30 kilometers 
south of the southernmost previously known to Cody (1996). 


Acknowledgments 

We thank Gerald A. Mulligan for the identification 
of Brassicaceae (Cruciferae) specimens; George Argus 
for the identification of Salix niphoclada; Stephen 
Darbyshire for identification of Agrostis humilis, Fes- 
tuca brachyphylla, Bromus japonicus, and Rumex mar- 
itimus ssp. fheginus; M. Barkworth for identification 
of Psathyrostachys juncea; Paul Catling for identifi- 
cation of Carex lasincarpa; Julie A. Dragon for iden- 
tification of Carex lenticularis var. dolia; R. Elven 
and H. Solstad for the identification of Papaver radi- 
catum ssp. kluanensis, R. Elven for the identification 
of Stellaria umbellata; A. A. Reznicek for identifica- 
tion of several species of Carex; M. Costea for iden- 
tification of Polygonum achoreum; Paul Catling for 
reviewing an earlier version of this manuscript and 
Leslie Cody for the many hours inputting this infor- 
mation on her computer. 


Literature Cited 

Britton, N. L., and P. A. Rydberg. 1901. Contributions to the 
botany of the Yukon Territory. 4. An enumeration of the 
flowering plants collected by R. S. Williams and J. B. Tarle- 
ton. Bulletin of the New York Botanical Garden 2: 149-187. 

Calder, J. A., and D. B. O. Savile. 1960. Studies in Saxifra- 
gaceae III. Saxifraga odontoloma and lyallii, and North 
American subspecies of S. punctata. Canadian Journal of 
Botany 38: 409-435. 

Cody, W. J. 1994. The flora of the Yukon Territory: Addi- 
tions, Range Extensions and Comments. Canadian Field- 
Naturalist 108: 428-476. 

Cody, W. J. 1996. Flora of the Yukon Territory. National 
Research Council (NRC) Press, Ottawa, Ontario, Canada. 
643 pages. 

Cody, W. J., C. E. Kennedy, and B. A. Bennett. 1998. New 
records of vascular plants in the Yukon Territory. Canadian 
Field-Naturalist 112: 289-328. 

Cody, W. J., C. E. Kennedy, and B. A. Bennett. 2000. 
New records of vascular plants in the Yukon Territory II. 
Canadian Field-Naturalist 114: 417-443. 

Cody, W. J., C. E. Kennedy, and B. A. Bennett. 2001. 
New records of vascular plants in the Yukon Territory II. 
Canadian Field-Naturalist 115: 301-322. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Cody, W. J., C. E. Kennedy, B. A. Bennett, and P. Caswell. 
2004. New records of vascular plants in the Yukon Terri- 
tory VI. Canadian Field-Naturalist 118: 558-578. 

Cody, W. J., C. E. Kennedy, B. A. Bennett, and V. Loewen. 
2002. New records of vascular plants in the Yukon Terri- 
tory IV. Canadian Field-Naturalist 116: 446-474. 

Cody, W. J., C. E. Kennedy, B. A. Bennett, and J. Stani- 
forth. 2003. New records of vascular plants in the Yukon 
Territory V. Canadian Field-Naturalist 117: 278-301. 

Cody, W. J., K. L. Reading, and J. M. Line. 1993. Addi- 
tions and range extensions to the vascular plant flora of 
the Continental Northwest Territories and Nunavut, Cana- 
da II. Canadian Field-Naturalist 117(3): 448-465. 

Douglas, G. W., and G. Ruyle-Douglas. 1978. Contributions 
to the floras of British Columbia and the Yukon Territory 
1, Canadian Journal of Botany 56: 2296-2302. 

Douglas, G. W., G. W. Argus, H. L. Dickson, and D. F. 
Brunton. 1981. The rare vascular plants of the Yukon. 
Syllogeus 28: 1-96. 

Douglas, G. W., D. Meidinger, and J. Pojar. 1998-2001. 
Illustrated flora of British Columbia. Volumes 1-6. 

Egger, M. 1992. Yukon Territory, Castilleja miniata Douglas 
ex. Hook. (Scrophulariaceae). Madrono 39: 244. 

Hultén, E. 1940. History of botanical exploration in Alaska 
and Yukon territories from the time of their discovery to 
1940. Botaniska Notiser 1940: 289-346. 

Hultén, E. 1968. Flora of Alaska and neighboring territo- 
ries. Stanford University Press, Stanford, California. 

Kartesz, J. T., and C. A. Meacham. 1999. Synthesis of the 
North American Flora Version 1.0. North Carolina Botan- 
ical Garden, Chapel Hill, North Carolina. 

Murray, D. F. 1995. New names in Papaver Section Mec- 
onella (Papaveraceae), Novon 5: 294-295. 

Porsild, A. E., and W. J. Cody. 1980. Vascular plants of 
continental Northwest Territories, Canada. National Muse- 
um of Natural Sciences, Ottawa, Ontario. 667 pages. 

Reznicek, A. A. 2002. 26}. Carex Linnaeus sect. Ammoglo- 
drin Dumortier, Fl. Belg., 146. 1827. Pages 306-307 in 
Flora of North American, North of Mexico. Edited by The 
Flora of North America Editorial Committee. Volume 23, 
Magnoliophyta: Commelinidae (inpart): Cyperaceae. Ox- 
ford University Press, New York. 

Standley, L. A., J. Cayouette, and L. Bruederle. 2002. 26r. 
Carex Linnaeus sect. Phacocyslis Dumortier, Fl. Belg., 
146. 1827. Pages 379-401 in Flora of North America, 
north of Mexico. Edited by The Flora of North America 
Editorial Committee. Volume 23, Magnoliophyta: Com- 
melinidae (in part): Cyperaceae. Oxford University Press, 
New York. 

Utech, F. H. 2002. Pages 50-409 in Liliaceae. Flora North 
America, north of Mexico. Edited by The Flora of North 
America Editorial Committee. Volume 26: 50-409. 

Welsh, S. L. 1974. Anderson’s Flora of Alaska and Adja- 
cent Parts of Canada. Brigham Young University Press, 
Provo, Utah, USA. 727 pages. 

Welsh, S. L. 2001. Revision of North American species 
Oxytropis, de Candolle “Leguminose”. Brigham Young 
University Press, Provo, Utah, USA 101 pages. 


Received 5 May 2005 
Accepted 23 July 2005 


Additions to the Flora of the Continental Northwest Territories from 
the Great Slave Lake Area 


PAUL M. CATLING 


Biodiversity, National Program on Environmental Health, Agriculture and Agri-food Canada, Wm. Saunders Bldg., Central 
Experimental Farm, Ottawa, Ontario K1A 0C6 Canada; e-mail: catlingp @agr.gc.ca 

Catling, Paul M. 2005. Additions to the flora of the continental Northwest Territories from the Great Slave Lake area. Canadian 
Field-Naturalist 119(3): 437-440. 


Eleven species are reported as new to the flora of the continental Northwest Territories. The new native species include 
Artemesia dracunculus, Chenopodium leptophyllum, Eleocharis erythropoda, Panicum capillare, Schoenoplectus pungens 
and Symphyotrichum lanceolatum subsp. hesperium var. hesperium. New alien species reported include Achillea ptarmica, 
Chaenorhinum minus, Galium aparine, Malva neglecta and Silene cserei. Sonchus arvensis, previously reported, is based 
on material referable to a subsp. uliginosus. Forms new to the flora include Achillea millefolium f. rosea and Actaea rubra f. 
neglecta. Locations, habitats and distinctive features are provided for the additional taxa. The Hay River lowland ecoregion 
is a floristically rich area that deserves more botanical exploration. 


Key Words: Additions, range extensions, vascular plants, Northwest Territories, Canada. 


The flora of the Northwest Territories (NWT) has 
been extensively studied by many botanists as outlined 
by Porsild and Cody (1980) with more recent work 
indicated by Catling et al. (2005*). Four days of study- 
ing the flora in southern NWT yielded 11 species not 
previously recorded. This suggests that either the flora 
is changing or that parts of the area are understudied, 
or both. These possibilities are briefly discussed and 
the additional species are included in an annotated 
list presented below. 

The latitude, longitude, and collection dates for all 
locations mentioned in the text are indicated in Table | 
and the coordinates are correct to 200 m. In all cases 


Phytogeographic aspects 
Native species 

All but one (Chenopodium leptophyllum) of the addi- 
tional native species are southern plants now reaching 
their northern limits at Hay River. This region south and 
west of Great Slave Lake is already known to be a dis- 
tinctive phytogeographic zone where many southern 
species reach their northern limits and is also an area 
where many arctic and alpine plants are absent but oth- 
erwise occur throughout NWT (e.g., Raup 1947, page 
66 lower figure). The area has been designated as the 
Hay River lowland ecoregion (number 64) of the Taiga 
Plain ecozone (Ecological Stratification Working Group 


the collector is P. M. Catling. Specimens documenting 
occurrences are preserved in the Agriculture and Agri- 
Food Canada collection (DAO, see Holmgren et al. 
1990 for an explanation of herbarium acronyms) in 
Ottawa. 


1995). It was designated as the Southern Boreal 
Province by McJannet et al. (1995). 

The climate warming trend in northern Canada will 
make it possible for plants to extend their distributions 


TABLE |. Locations, habitats, collection dates and latitude and longitude of places in Northwest Territories where additions 


to the flora were discovered by P. M. Catling in 2003. 


Habitat and Location Date Latitude Longitude 
periodically flooded shore 30 July 2003 60.4900 115.7304 
of Hay River at Hay River 

open vacant land and shoreline 30 July 2003 60.8214 115.7304 
at port of Hay River 

bank of Hay River near West 30 July 2003 60.8536 115.7743 
Channel bridge N side of Hay River 

bulldozed area beside Niven 30 July 2003 62.4603 114.3745 
Lake, N side of Yellowknife 

vacant lots in Yellowknife 30 July 2003 62.4500 114.4300 

open sand prairie, Taltson River 23 July 2003 60.3554 111.2760 
W of Fort Smith 

open rocky ridge, Ingraham Trail, 28 July 2003 62.5037 114.2764 


E of Yellowknife 


437 


438 


northward but warming is not likely to be an explana- 
tion for new records of native species reported here. 
Warming to the extent of promoting northward range 
extension may not have occurred yet and the new native 
species are easily overlooked in cursory botanical 
inventory. Consequently they are likely to be long- 
established in the area. Unlike the Fort Simpson, Fort 
Smith and Wood Buffalo areas in the Hay River low- 
land, the region of Hay River itself has not been suffi- 
ciently studied from a botanical viewpoint. There are 
only few and small collections from the area including 
those of W. H. Lewis in 1951 (at DAO) reported by 
Cody (1956) and collections made in 1971 by L. 
Dahike (at CAN and WAT) and by P. Ducruc, both 
included in Porsild and Cody (1980). An examination 
of the maps in Porsild and Cody (1980) indicates that 
many plants likely to occur in the area have not been 
recorded. 


Introduced species 

There are few published studies of the flora of man- 
made disturbed habitats such as the Norman Wells 
pipeline study (Cody et al. 2000). In an unpublished 
report involving the non-cultivated urban flora of Yel- 
lowknife, Steinecke (2001*) gathered information on 
135 sites within the city limits. The sampled habitats 
included roadsides, lawns and vacant lots. A total of 142 
species were recorded in the town, 55% of which were 
alien. Of these, 25 are additions to the flora (Porsild 
and Cody 1980; Catling et al. 2005*), including Acer 
negundo L., Aconitum sp., Alopecurus occidentalis 
Scribn. & Tweedy (probably A. arundinaceus Sobol 
which is not new), Amsinckia menziesii (Lehm.) Nels. 
& Macbr., Artemesia absinthium L., Campanula rapun- 
culoides L., Dianthus barbatus L., Halenia deflexa 
(Smith) Giseb., Hordeum vulgare L., Malva rotundi- 
folia L., Panicum miliaceum L. (introduced with bird 
seed around feeders but possibly not persisting), Por- 
tulaca oleracea L., Ranunculus repens L., Rheum rha- 
barbarum L., Rumex crispus L., Setaria glauca (L.) 
Beauv., Sonchus uliginosus Bieb., Spiraea cf. betuli- 
folia Pallas and Tropaeolum sp. Since this interesting 
study is unpublished and there are no specimen vouch- 
ers, the records are not accepted as additions, but they 
are nevertheless of interest and, hopefully, they will 
be supported by vouchers in the future. 

As settlement expands and road traffic increases it 
is likely that more introduced species will expand into 
the region by gradual spread along corridors of dis- 
turbed habitats such as roads, but also through long 
distance dispersal by people incidentally transporting 
soil and seeds on or in vehicles. That such spread of 
alien species is continuing is suggested by the addition- 
al species reported by Steinecke (200*, listed above) 
and the five additional alien species reported here. 
The latter originated from open disturbed sites in either 
the town of Yellowknife or the town of Hay River. 
More alien species are expected to enter the region 
and some may begin to impact native flora and fauna, 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


but with the possible exception of Phalaris canariensis 
L. (recently reported to be a local problem), there is 
little evidence of spread into native plant communities. 


Annotated List of Additional Species 
Achillea millefolium L. f. rosea Rand. & Redf. 
YARROW (rose-flowered form) 

Although the species is well known from NWT, this 
unusual form with deep rose-purple (instead of white) 
ligules has not been recorded previously. Found in a 
bulldozed area beside Niven Lake on N side of Yel- 
lowknife, it is occasionally cultivated and likely a gar- 
den escape. 


Achillea ptarmica L. 
PEARL YARROW 

Found in an open bulldozed area beside Niven Lake 
on N side of Yellowknife, this introduced species differs 
from A. sibirica Ledeb. in its serrated instead of pecti- 
nate leaves and longer ligules to 5 mm. This species is 
sometimes introduced with “wildflower” seed mixtures. 


Actaea rubra (Ait.) Willd. f. neglecta (Gillman) Robins. 
RED BANEBERRY (white berry form) 

This white-fruited form of this native plant 1s abun- 
dant in some parts of the extensive North American 
range, but previously only “bright red or rarely pink” 
fruits have been reported for NWT (Porsild and Cody 
1980). Several plants were found on a wooded bank of 
the Hay River near bridge on north side of town. 


Artemesia dracunculus L. 
DRAGON WORMWOOD 

This native, linear-leaved species of dry hills and 
prairies was expected to be discovered in NWT by 
Porsild and Cody (1980). It was found in open sandy 
ground on banks of the Hay River at Hay River. 


Chaenorhinum minus (L.) Lange 
DWARF SNAP-DRAGON 

This distinctive, glandular-hairy, introduced plant 
was frequent in open gravelly areas and along the rail- 
way at the port of Hay River. The 5-7 mm flowers are 
bluish-purple with yellow in the throat. 


Chenopodium leptophyllum (Moquin-Tandon) Nuttall 
ex S. Watson 
NARROW-LEAF GOOSEFOOT 

Porsild and Cody (1980) listed this species but ex- 
cluded it based on the Richardson specimen collected at 
Fort Franklin on Great Bear Lake “where it was surely 
an ephemeral introduction.” Clements and Mosyakin 
(2003) show its occurrence in Alberta near the NWT 
border but not in NWT. It was found on a disturbed 
sandy portion of an open prairie slope adjacent to the 
Taltson River east of Fort Smith. At this location it 
occurred around an open sandy blowout on dry, open, 
south-facing slopes dominated by Carex siccata Dewey 
with Calamagrostis purpurascens R. Brown and many 
other native species of dry, open ground, including 
Agrostis scabra Willdenow, Arabis holboellii Horne- 


2005 


man, Anemone multifida Poiret and Pinus banksiana 
Lamb. It was also found on top of a rocky ridge E of 
Yellowknife. This was also a natural, open and rather 
dry plant community dominated by native species, 
including Androsace septentionalis L., Arctostaphylos 
uva-ursi (L.) Spreng., Artemesia hyperborea Rydb., 
Calamagrostis purpurascens R. Br., Carex supina 
Willd., Juniperus communis L., Juniperus horizontalis 
Moench, Rosa acicularis Lindl., Saxifraga tricuspidata 
Ait. and Senecio pauperculus Michx. The flora was 
entirely native and had not been disturbed by man. 
Consequently Chenopodium leptophyllum is consid- 
ered native. 


Eleocharis erythropoda Steudel 
BALD SPIKE-RUSH 

Frequent on the periodically flooded shore of the 
Hay River at Hay River, this native plant was reported 
for the Northwest Territories by Smith et al. (2002) but 
accidentally omitted from the recent compilation of 
additions (Catling et al. 2005*) so included here. It dif- 
fers from E. palustris (L.) Roemer & Schultes by the 
basal scale completely or nearly surrounding the stem 
(instead of encircling only 2/3) and relatively narrow 
stems 0.3-1.4 mm thick (instead of 0.5-5 mm thick). 
See under “Panicum capillare” for associated species. 


Galium aparine L. 
CLEAVERS 

Found in vacant lots and disturbed open places in 
Yellowknife, this species differs from G. triflorum by 
the retrorse barbs on leaves and stems. There are possi- 
bly both native and introduced races; the plant of open 
disturbed sites is probably introduced. 


Malva neglecta Wallr. 
DWARF MALLOw 

A few plants of this introduced species were found in 
vacant lots in Yellowknife. This is the first report of a 
species of Malva in NWT. The rounded leaves and axil- 
lary flowers with petals twice as long as the sepals are 
distinctive. 


Panicum capillare L. var. occidentale Rydberg 
COMMON PANIC GRASS 

This is the first report of a species in this genus for 
NWT. The exerted panicles and long-acuminate spike- 
lets, many with pedicels shorter than the spikelet, sug- 
gested var. occidentale. The plants were sporadic on 
the periodically flooded shore of the Hay River at Hay 
River. A native species, it often occurs in disturbed 
sites and is sometimes thought to be an introduction, 
but on the periodically flooded shoreline it occurred 
with a diversity of native species in one of its charac- 
teristic natural habitats. Close plant associates includ- 
ed Aster modestus Lindl., Caltha natans Pall., Carex 
sychnocephala Carey, Cicuta maculata var. angustifolia 
Hook., Deschampsia caespitosa (L.) Beauv., Eleocharis 
erythropoda Steud., Galium triflorum Michx., Gna- 
phalium uliginosum L., Juncus nodosus L., Limosella 
aquatica L., Poa palustris L., Potentilla anserina L., 


CATLING: ADDITIONS TO THE FLORA OF THE NORTHWEST TERRITORIES 


439 


Ranunculus scleratus L., Ranunculus macounii Britt., 
Rorippa palustris (L.) Besser, Rumex maritimus L., 
Schoenoplectus pungens (Vahl) Palla, Scirpus micro- 
carpus Presl and Spartina gracilis Trin. 


Schoenoplectus pungens (Vahl) Palla 
(Scirpus americanus auct. non Persoon) 
COMMON THREE-SQUARE BULRISH 

This native plant was frequent along the periodically 
flooded shore of Hay River at Hay River. It is distinctive 
in the lateral (instead of terminal) inflorescence with 
stemless spikelets and the sharply three-angled stems. 
This occurrence may be an extension of known range of 
approximately 700 km based on Packer’s (1983) map. 
See under “Panicum capillare” for associated species. 


Silene cserei Baumg. 
BALKAN CATCHFLY 

Found along the railway and in adjacent open sandy 
soil at port of Hay River, this tall introduced species 
(2-8 dm) has 3 styles, lacks glutinous bands on the stem 
and is without an inflated calyx. 


Symphyotrichum lanceolatum (Willd.) G. L. Nesom 
subsp. hesperium (Gray) Nesom var. hesperium (Aster 
lanceolatus Willd. subsp. hesperius (A. Gray) Semple 
& Chmielewski, Aster hesperius A. Gray) 
WESTERN WILLOW ASTER 

This native plant was found on the periodically flood- 
ed shore of Hay River at Hay Riverand at the port of 
Hay River. At least 100 plants were seen and they had 
either white or blue ray flowers. This species has green 
leaves 5-15 mm wide instead of purplish-green leaves 
less than 5 mm wide as in A. borealis (T. & G.) Prov. 
(previously A. junciformis Rydberg). Unlike A. spathu- 
latus Lindley, the stems are pubescent. This species is 
well known from the Lake Athabasca area of northern 
Alberta. 


Additional Notes 

Sonchus arvensis L. subsp. uliginosus (Bieb.) Nyman 
(S. arvensis var. glabrescens Guenth., Grab. & Wimmer, 
S. uliginosus Bieb.) 

This introduced species is frequent in disturbed sites 
at Hay River and Yellowknife, but is not an addition 
because the inclusion of “Sonchus arvenvis L.” in Por- 
sild and Cody (1980) is not based on material of subsp. 
arvensis but rather on material referable to this variety 
(based on examination of specimens collected at Hay 
River (Cody 14839, DAO) and on the Mackenzie high- 
way (Thieret 6172, DAO). 


Acknowledgments 

Suzanne Carriere, Ecosystem Management Biolo- 
gist, Wildlife & Fisheries Division, Department of 
Resources, Wildlife & Economic Development, pro- 
vided extensive help and support. Bill Cody, Curator 
Emeritus, with AAFC in Ottawa provided essential 
information and comments on the manuscript. Deb- 
orah Johnson and Mike Fournier assisted with plant 
collecting. 


440 


Documents Cited (marked * in text) 

Catling, P. M., W. J. Cody, and G. Mitrow. 2005. A compila- 
tion of additions to the flora of the continental portions of 
Northwest Territories and Nunavut. Botanical Electronic 
News 353. www.ou.edu/cas/botany-micro/ben/ben353.html. 

Steinecke, K. 2001. Urban flora and plant communities in sub- 
arctic settlements: two case examples from Yellowknife, 
Canada, and Reykjavik, Iceland. University of Bremen, 
Bremen, Germany. 34 pages. 


Literature Cited 

Clements, S. E., and S. L. Mosyakin. 2003. Chenopodium 
Linnaeus. Pages 275-299 in Flora of North America Edi- 
torial Committee, eds. Flora of North America. Volume 4, 
Magnoliophyta: Caryophyllidae, part 1. Oxford University 
Press, New York. 

Cody, W. J. 1956. New plant records for northern Alberta and 
southern Mackenzie District. Canadian Field-Naturalist 
70: 101-130. 

Cody, W. J., K. L. MacInnes, J. Cayouette, and S. Dar- 
byshire. 2000. Alien and invasive native vascular plants 
along the Norman Wells Pipeline, District of Mackenzie, 
Northwest Territories. Canadian Field-Naturalist 114: 126- 
137. 

Ecological Stratification Working Group. 1995. A National 
Ecological Framework for Canada. Agriculture and Agri- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Food Canada, Research Branch, Centre for Land and Bio- 
logical Resources Research and Environment Canada, State 
of the Environment Directorate, Ecozone Analysis Branch, 
Ottawa. Report and National Map at 1:7 500 000 scale. 

Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. 
Index herbariorum, part 1. The herbaria of the world. 
New York Botanical Garden. Bronx, New York. 693 pages. 

McJannet, C. L., G. W. Argus and W. J. Cody. 1995. Rare 
vascular plants in the Northwest Territories. Canadian 
Museum of Nature Syllogeus (73). 104 pages. 

Packer, J. G. 1983. Flora of Alberta. Second edition, revised. 
University of Toronto Press, Toronto. 687 pages. 

Porsild, A. E., and W. J. Cody. 1980. Vascular plants of 
continental Northwest Territories. National Museums of 
Canada, Ottawa. 667 pages. 

Raup, H. M. 1947. The botany of southwestern Mackenzie. 
Sargentia 6: 1-275. 

Smith S. G., J. J. Bruhl, M. S. Gonzalez-Elizondo, and F. 
J. Menapace. 2002. 7. Eleocharis. Pages 60-120 in Flora 
of North America Volume 23, Magnoliophyta: Com- 
melinidae, (in part): Cyperaceae. Edited by Flora of North 
America Editorial Committee, Oxford University Press, 
New York. 


Received 31 March 2005 
Accepted | September 2005 


Notes 


Possible Tool Use by Beavers, Castor canadensis, in a Northern Ontario 
Watershed 


D. M. BARNES 


Joint Appointment in Department of Biology and Faculty of Forestry and the Forest Environment, Lakehead University, 955 
Oliver Road, Thunder Bay, Ontario P7B 5E1 Canada 


Barnes, D. M. 2005. Possible tool use by Beavers, Castor canadensis, in a northern Ontario watershed. Canadian Field-Nat- 
uralist 119(3): 441-443. 


At a remote active Beaver (Castor canadensis) dam site in the Chapleau Crown Game Preserve of northern Ontario, I noted 
an instance where a Beaver had used a willow stem as a prop to allow it to cut other willow stems at an unrealistic height. 
The use of this tool (prop) meant less land foraging that in turn reduced the risk of predation and prevented undue thermal 
stress. This is the first account in the literature of a Beaver possibly using a tool to aid in foraging. 


Key Words: Beaver, Castor canadensis, boreal forest, foraging, tool use, Ontario. 


Tool use has long fascinated behavioural scientists 
involved in understanding animal evolution (Washburn 
1960; Lancaster 1968). Over the years, researchers have 
documented the use of tools in insects (Wilson 1975), 
birds (Lack 1947; Chisholm 1954; van Lawick-Goodall 
and van Lawick-Goodall 1966; Morse 1968; van Law- 
ick-Goodall 1970; Anderson 1989; Marks and Hall 
1992), and mammals (Kortland 1962; van Lawick- 
Goodall 1968; van Lawick-Goodall 1970; Chevalier- 
Skolnikoff and Liska 1993; Hart and Hart 1994; Toki- 
da et al. 1994). 

Scientists have recognized that Beavers (Castor can- 
adensis) have the ability to construct elaborate dams, 
lodges, and canals. Despite these achievements, there 
has been only one example of tool use documented in 
the literature. While studying two captive Beavers at the 
Berne Brain Anatomy Institute, Pilleri (1983) docu- 
mented an ingenious example of tool use. He kept 
Beavers in a concrete pool that had a constant supply 
of fresh water. The overflow from the pool went into a 
vertical metal pipe via three 0.8 cm holes. From a sup- 
ply of twigs left for their use, the Beavers successfully 
cut three sticks to exact dimensions and wedged them 
into the three outflow holes completely stopping the 
flow of water. 

In 1993, while conducting research on Beavers 
(Barnes and Mallik 1996, 1997, 2001), I documented 
another possible example of Beaver tool use. At a re- } bs Feet 
mote active Beaver dam site within the Swanson River "=m a hel ed AE RE. e- 
drainage area of the Chapleau Crown Game Preserve FIGURE 1. A photograph showing the Beaver’s use of a willow 
(CCGP), 48°04'N, 83°15'W of northern Ontario (Barnes stem as a prop to enable cutting at the elevated height 
and Mallik 2001), I found a willow (Salix spp.) clump SOE eS ene Wee ert ees 
which had stems cut at a height of approximately 1 m lighter-coloured stem supporting the camera lens cap 


; ie (diameter — 6 cm). Note that the Beaver has cut both 
above the ground (Figure |). This is an extraordinary the upper and lower ends of the prop; the lower end 


height considering that Beavers, on average, cut at an cut is hard to see due to ground cover. Unfortunately, 
average height of 30 cm above the ground (Johnston the photograph does not show the 45° leaning angle. 


44] 


442 


and Naiman 1990; Barnes and Mallik 1997). I made 
a careful examination of the area and found that there 
was no apparent way that Beavers could have cut the 
stems at such a height. When I studied the willow 
clump more closely, I noted that there was a freshly 
cut willow stem, approximately 12 cm in diameter, 
leaning against the main stem of the willow clump; 
its approximate angle was 45° (Figure 1). In addition, 
I observed cutting at both ends of the leaning willow 
segment. I first thought that the stem had fallen into 
place. However, this explanation does not seem pos- 
sible for three reasons. First, the willow stem would 
have to have fallen from the overhead clump and 
lodged in a leaning position. When one compared 
colour and texture of bark, the leaning willow seg- 
ment was clearly different from the stems of the wil- 
low clump (Figure 1). Second, even if we assume the 
willow segment came from the clump, it would have 
to fall from a height in order to land in the proper 
position. From Figure 1, cuts directly above the lean- 
ing willow are clearly too high for Beavers to harvest 
from ground level; i.e., approximately 1 m. Third, the 
fallen stem would have had to land exactly into the 
correct position; 1.e., 45° angle and distance of 45 cm 
below the cut stems of the willow clump (Figure 1). 

Another possible explanation as to why Beavers were 
able to cut stems at such a height may have some- 
thing to do with snow levels. Northern Beavers in the 
late winter and early spring will often forage over snow 
in order to obtain fresh food along impoundments 
and streams (Mech 1966; Peterson 1977). To gauge the 
reliability of this probable scenario, one must have a 
better appreciation for the general area. The dam itself 
was 118 m in length and a height of 1 m. The willow 
clump was located a distance of approximately 10 m 
downstream from the dam. I noted that the overflow 
from the dam created four smaller streams; the largest 
of these had an average width and depth of 1.6 m and 
0.17 m, respectively. In my opinion, if Beavers were to 
exit the waterway in winter, their most optimal strategy 
would have been to utilize the impoundment edge. 
Rue (1964) noted that in winter, water levels in Beaver 
impoundments often become reduced leaving air pock- 
ets along the shore, and these become convenient places 
for Beavers to access land. To travel overland to the 
willow clump the Beaver would have had to climb over 
the dam and wade through deep snow for 10 m, as the 
downstream water channels were too small to afford 
any great advantage. This choice would be highly un- 
realistic. Barnes and Mallik (2001) showed that in the 
ice-free seasons, these northern Beavers forage for food 
trees in close proximity to water. Their study showed 
that Beavers harvested 71% of Trembling Aspen (Pop- 
ulus tremuloides) stems within 10 m of the impound- 
ment edge. Beavers restricted their foraging range 
because of the risk of predation by Gray Wolves (Canis 
lupus) (Barnes and Mallik 2001). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


In response to this perplexing situation, I propose 
an alternative explanation. I believe that a Beaver 
deliberately cut and transported a willow stem segment 
and placed it against the main stem of the willow clump. 
This prop enabled the Beaver to establish an elevated 
foraging position that facilitated the cutting of nar- 
rower stems located above the thickened stems of the 
clump. However, this reasoning does not explain how 
Beavers were able to harvest some of the highest stems 
(Figure 1). Like Rue (1964), I have noticed some in- 
stances where Beavers have shown surprising climb- 
ing agility. I, therefore, believe that these Beavers may 
well have used this prop in conjunction with other 
clump features to establish elevated platforms from 
which to cut (Figure 1). 

This type of behaviour is characteristic of tool use 
as reported in the literature. In his treatise on the evo- 
lution of tool use in feeding animals, Alcock (1972) 
characterizes tool use as the manipulation of an inan- 
imate object that improves the organism’s efficiency 
in altering the position or form of some other object. 
Further, the organism should not be able to manufac- 
ture the object internally. Recently, Michener (2004) 
found that North American Badgers (TJaxidea taxus) 
facilitated the capture of Richardson’s Ground Squirrel 
(Spermophilus richardsonii) by using soil to plug their 
tunnels. In like manner, by using the cut stem as a prop, 
the Beaver was able to gain a better position to cut the 
higher positioned narrow willow stems. Clearly, the 
propped stem afforded the Beaver a platform with an 
optimal foraging height. 

This observation is significant for two reasons: (1) 
this account represents the first documented case of 
Beavers using a tool to facilitate foraging activity; 
and (2) the use of a tool, from an optimal foraging 
perspective, provided the Beaver with an ecological 
advantage by reducing the foraging time on land. In 
boreal situations, such as the CCGP, a lessening of 
foraging time means reduced exposure to Gray Wolves. 
Potvin et al. (1992) demonstrated that Gray Wolves 
are highly effective predators. In subsequent studies, 
Basey and Jenkins (1995) further showed that Beavers 
actually trade off maximum profitability against min- 
imization of Wolf predation. In addition, Beavers expe- 
rience thermal stress with prolonged exposure to over- 
land foraging activity (Barnes and Mallik 2001). 


Acknowledgments 

I acknowledge the financial assistance of Ducks 
Unlimited, the Ontario Ministry of Universities and Col- 
leges, and Environmental Youth Corps. I am indebted to 
A. Mallik, Department of Biology, Lakehead University 
and C. Todesco of the Ontario Ministry of Resources 
for their guidance and logistic support. I am grateful 
to F. DeGagne for his technical assistance during the 
field work. In addition, I thank B. Barnes, Department 
of Biology, Lakehead University for her proof-reading 
skills. 


2005 


Literature Cited 

Alcock, J. 1972. The evolution of the use of tools by feeding 
animals. Evolution 26(3): 464-473. 

Anderson, S. 1989. Tool use by the Fan-tailed Raven (Corvus 
rhipidurus). Condor 91: 999. 

Barnes, D. M., and A. U. Mallik. 1996. Use of woody plants 
in construction of beaver dams in northern Ontario. Cana- 
dian Journal of Zoology 74: 1781-1786. 

Barnes, D. M., and A. U. Mallik. 1997. Habitats influencing 
Beaver dam establishment in a northern Ontario watershed. 
Journal of Wildlife Management 61: 1371-1377. 

Barnes, D. M., and A. U. Mallik. 2001. Effects of Beaver, 
Castor canadensis, herbivory on streamside vegetation in 
a northern Ontario watershed. Canadian Field-Naturalist 
115: 8-21. 

Basey, J. M., and S. H. Jenkins. 1995. Influences of preda- 
tion risk and energy maximization on food selection by 
beavers (Castor canadensis). Canadian Journal of Zoology 
73: 2197-2208. 

Chevalier-Skolnikoff, S., and J. Liska. 1993. Tool use by 
wild and captive elephants. Animal Behaviour 46: 209-219. 

Chisholm, A. H. 1954. The use by birds of “tools” or “in- 
struments”. Ibis 96: 380-383. 

Hart, B. L., and L. A. Hart. 1994. Fly switching by Asian 
elephants: tool use to control parasites. Animal Behaviour 
48: 35-45. 

Johnston, C. A., and R. J. Naiman. 1990. Browse selection 
by beaver: effects on riparian composition. Journal of Forest 
Research 20: 1036-1043. 

Kortland, A. 1962. Chimpanzees in the wild. Scientific Amer- 
ican 206: 128-138. 

Lack, D. 1947. Darwin’s finches. Cambridge University Press, 
Cambridge, England. 

Lancaster, J. 1968. On the evolution of tool-using behaviour. 
American Anthropology 70: 55-66. 

Marks, J. S., and C. S. Hall. 1992. Tool use by Bristle-thighed 
curlews feeding on Albatross eggs. Condor 94: 1032-1034. 


NOTES 


443 


Mech, L. D. 1966. The Wolves of Isle Royale. Fauna of the Na- 
tional Parks of the United States Fauna Series 7. 210 pages. 

Michener, G. R. 2004. Hunting techniques and tool use by 
North American badgers preying on Richardson’s ground 
squirrels. Journal of Mammalogy 85: 1019-1027. 

Morse, D. H. 1968. The use of tools by Brown-headed 
Nuthatches. Wilson Bulletin 80: 220-224. 

Peterson, R. O. 1977. Wolf ecology and prey relationships 
on Isle Royale. National Park Service Scientific Monograph 
Series Number 11. 210 pages. 

Pilleri, G. 1983. Ingenious tool use by the Canadian Beaver 
(Castor canadensis) in captivity, Pages 99-102 in Investi- 
gations on Beavers, Volume |. Edited by G. Pilleri. Brain 
Anatomy Institute, Berne, Switzerland. 

Potvin, F., L. Breton, and M. MacQuart. 1992. Impact of an 
experimental wolf reduction on beaver in Papineau-LaBelle 
Reserve, Quebec. Canadian Journal of Zoology 70: 180-183. 

Rue, L. L., HI. 1964. The world of the Beaver. J. B. Lip- 
pincott Co., Philadelphia. 158 pages. 

Tokida, E., I. Tanaka, H. Takefushi, and T. Hagiwara. 1994. 
Tool-using in Japanese macaques: use of stones to obtain 
fruit from a pipe. Animal Behaviour 47: 1023-1030. 

van Lawick-Goodall, J. 1968. The behaviour of free living 
chimpanzees in the Gambe Stream Preserve. Animal Be- 
haviour Monograph |: 163-311. 

van Lawick-Goodall, J. 1970. Tool-using in primates and 
other vertebrates. Advances in the Study of Behaviour 3: 
195-249. 

van Lawick-Goodall, J., and H. van Lawick-Goodall. 1966. 
Use of tools by the Egyptian Vulture, Neophron percnop- 
terus. Nature 212: 1468-1469. 

Washburn, S. L. 1960. Tools and human evolution. Scientific 
American 203: 63-75. 

Wilson, E. O. 1975. Sociobiology — The new synthesis. The 
Belknap Press of Harvard University Press, Cambridge, 
Massachusetts. 697 pages. 


Received 1 October 2002 
Accepted 30 June 2005 


Predation of a Bat by American Crows, Corvus brachyrhynchos 


KARA L. LEFEVRE 


Department of Zoology, University of Toronto, 25 Harbord Street, Toronto, Ontario M5S 3G5 Canada; email: k.lefevre@ 


utoronto.ca 


Lefevre, Kara L. 2005. Predation of a bat by American Crows, Corvus brachyrhynchos. Canadian Field-Naturalist 119(3): 


443-444. 


This report documents predation and possible group hunting of a bat by American Crows (Corvus brachyrhynchos). | 
observed a group of several crows appear to cooperate in order to ground and catch a bat. Predation of bats by birds in 
North America is relatively rare, and bat predation by crows has not been previously reported. A cooperative attack would 
be consistent with other group behaviours of the species, including territoriality, breeding and foraging. 


Key Words: American Crow, Corvus brachyrhynchos, bats, Chiroptera, Myotis lucifugus, predation, Ontario. 


Several bird species are known to feed occasionally 
on bats—mainly birds of prey, but also members of 
families Laridae and Corvidae, among others (Speak- 
man 1991; Radzicki et al. 1999; Lee and Kuo 2001). 
The omnivorous American Crow (Corvus brachyrhyn- 
chos) is one of North America’s most widespread birds 


(Verbeek and Caffrey 2002). This opportunistic species 
is known as both a scavenger of anything edible and a 
predator of diverse taxa, including small mammals 
(e.g., Nocera 2000; Verbeek and Caffrey 2002). To my 
knowledge the incident reported here is the first pub- 
lished record of bat predation by C. brachyrhynchos. 


444 


I observed an episode of bat predation by American 
Crows along the Rideau River in Ottawa, Ontario 
(45°19'N, 75°40'W) on 18 April 1996 at 17:45 EST. 
Visibility was excellent due to the absence of foliage 
following winter. I noticed a group of eight crows 
perched approximately 10 m high, at the edge of a 
deciduous woodlot beside the river. The woods bor- 
dered on an open field of dead grass. Four of the eight 
crows remained perched while the rest became 
involved in a chase of a small bat (Order Chiroptera) 
about the size of a sparrow. Based on size and relative 
species abundances it was possibly a Little Brown 
Bat (Myotis lucifugus), but seven -bat species that 
occur in the area fit the size description (A. J. Erskine 
and S. Peters, personal communication). Mid-April is 
early in the season for bat activity in eastern Ontario: 
M. lucifugus females leave the hibernacula at this time 
and fly to the nurseries, while males remain torpid 
until mid-May (Gerson 1984). 

During the chase the bat flew jerkily in different 
directions, approximately 5—7 m above the ground, 
eluding capture by 1-3 crows that chased it at any 
given time. Each time crows approached closely the 
bat dropped in altitude, effectively evading them. The 
crows attempted to use their bills but not their feet to 
strike the bat. The bat seemed to escape after three 
minutes of this activity, and the crows stopped chase 
for a brief time. One crow then darted after the bat 
and two others followed. Two of the three crows 
moved in and appeared to “sandwich” the bat about 3 
m above the ground. They likely struck the bat 
because it dropped suddenly to the ground, with the 
third crow in pursuit. The three crows immediately 
moved to the ground: one struck the bat hard enough 
to make it pop up into the air above their heads, and 
then picked the bat up in its bill. There was no appar- 
ent dispute for possession of the bat. The three crows 
flew up to where the others were perched; the captor 
and another of the chasers landed in the same branch, 
and then the captor flew approximately 50 m South 
along the edge of the words to a new perch, also 10 m 
above the ground. The bat was still in its bill and none 
of the other crows followed. The captor attempted awk- 
wardly to bite/peck at the bat held in its feet. After one 
minute at this new perch, the captor flew beyond view 
with the bat in its bill. The other crows remained in 
the original area, some perched in the trees and oth- 
ers in the field near where the bat had been grounded. 

These observations may represent an episode of co- 
operative hunting. Hendricks and Schlang (1998) re- 
ported similar cooperative aerial attacks by the con- 
generic Common Raven (C. corax) on adult pigeons. 
Such group hunting by C. brachyrhynchos would be 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


consistent with adaptive cooperation observed in this 
species including group territoriality (Caccamise et al. 
1997), cooperative breeding (Caffrey 1992), and group 
foraging (Kilham 1989; Verbeek and Caffrey 2002). 
The absence of any apparent competition for the cap- 
tured bat suggests that the captor could have been an 
adult crow with the other participants being imma- 
ture. Alternatively, it is also possible that this was an 
instance of several crows attempting to hunt the same 
bat individually, without any real cooperation. 

This observation is noteworthy because direct pre- 
dation events are seldom observed, predation of bats by 
birds in North America is apparently rare, and be- 
cause this is the first account of American Crows hunt- 
ing bats. Based on a review of the impact of bird pre- 
dation on bat mortality (Speakman 1991), it is unlikely 
that crows pose any significant predation threat to 
North American bat populations. 


Acknowledgments 

I thank A. J. Erskine, Sandra Peters, and one anony- 
mous reviewer for comments that improved the manu- 
script. 


Literature Cited 

Caccamise, D. F., L. M. Reed, J. Romanowski, and P. C. 
Stouffer. 1997. Roosting behavior and group territoriality 
in American Crows. Auk 114: 628-637. 

Caffrey, C. 1992. Female-biased delayed dispersal and 
helping in American Crows. Auk 109: 609-619. 

Gerson, H. 1984. Habitat management guidelines for bats of 
Ontario. Ontario Ministry of Natural Resources. 

Hendricks, P., and S. Schlang. 1998. Aerial attacks by Com- 
mon Ravens, Corvus corax, on adult feral pigeons, Colum- 
bia livia. Canadian Field-Naturalist 112: 702-703. 

Kilham, L. 1989. The American Crow and the Common 
Raven. Texas A & M University Press, College Station. 

Lee, Y. F., and Y. M. Kuo. 2001. Predation on Mexican free- 
tailed bats by Peregrine Falcons and Red-tailed Hawks. 
Journal of Raptor Research 35: 115-123. 

Nocera, J. J. 2000. Predation on an Eastern Chipmunk, Jamias 
striatus, by an American Crow, Corvus brachyrhynchos. 
Canadian Field-Naturalist 114: 326-327. 

Radzicki, G., J. Hejduk, and J. Banbura. 1999. Tits (Parus 
major and Parus caeruleus) preying upon hibernating bats. 
Ornis Fennica 76: 93-94. 

Speakman, J. R. 1991. The impact of predation by birds on 
bat populations in the British Isles. Mammalian Review 
21: 123-142. 

Verbeek, N. A. M., and C. Caffrey. 2002. American Crow 
(Corvus brachyrhynchos) in The Birds of North America 
(647). Edited by A. Poole and F. Gill. The Birds of North 
America, Inc., Philadelphia, Pennsylvania. 


Received | July 2004 
Accepted 2 September 2005 


Mortality of Deer Mice, Peromyscus maniculatus, in Wire Mesh Live- 


Traps: A Cautionary Note 


Tuomas S. JuNG!, and KIERAN S. O’ DONOVAN? 


' Yukon Department of Environment, Box 2703, Whitehorse, Yukon Y1A 2C6 Canada 
2 308A Klukshu Boulevard, Whitehorse, Yukon Y1A 3Y1 Canada 


Jung, Thomas S., and Kieran S. O’ Donovan. 2005. Mortality of deer mice (Peromyscus maniculatus) in wire mesh live-traps: 
a cautionary note. Canadian Field-Naturalist 119(3): 445-446. 


Live-capture of animals occasionally results in the death of some individuals. Here, we report upon an unusual occurrence 
of trap-related mortality observed in Deer Mice (Peromyscus maniculatus) captured in wire mesh live-traps during field 
sampling in southeastern Yukon. Eight of 85 marked individuals (9.4%) were found with their snouts caught in the wire 
mesh of our live-traps; four of these individuals were found dead. We suggest a modification to Ugglan live-traps that would 


likely decrease such incidents. 


Key Words: mortality, Deer Mouse, Peromyscus maniculatus, Live-trapping, Yukon. 


Biologists often rely upon data obtained through 
the live-capture of individuals. Field sampling of ani- 
mal populations and communities, however, occasion- 
ally results in the death of some individuals. Mortality 
can compromise data collection when experimental 
designs require individuals to be live-captured, marked 
and, later, recaptured. When deaths are the result of 
equipment or technique then it is incumbent upon field 
researchers to share this information and seek means 
to reduce capture mortality, for both ethical and data 
collection reasons (e.g., Jung et al. 2002). Here, we 
report upon an unusual incident of trap-related mor- 
tality of Deer Mice (Peromyscus maniculatus) during 
field sampling in southeastern Yukon. 

As part of a study of small mammal communities in 
the boreal forest near Watson Lake, Yukon (60.06° N, 
128.70° W), we used Ugglan live-traps (Model 3 
Lemming Special, Granhab, Marieholm, Sweden) to 
live-capture small mammals during September 2003. 
Unlike some other commonly used types of small mam- 
mal live-traps like Sherman traps (H. B. Sherman Traps, 
Tallahassee, FL) or Longworth traps (Longworth Sci- 
entific Instruments Co., United Kingdom), Ugglan 
traps are constructed of wire mesh and the trap door 
is elevated and gravity controlled (as opposed to spring 
loaded). Our traps measured 250 x 78 x 65 mm and 
had a 6 X 6 mm wire mesh around the top and sides of 
the trap. A weather shield made of sheet metal covered 
most of the wire mesh. 

We captured 85 individual Deer Mice 167 times dur- 
ing our study. Eight individuals (9.4% of individuals 
captured) were found with their snouts caught in the 
wire mesh of the traps (Figure |). Because their upper 
incisors were through to the other side of the mesh, 
they were unable to free their snouts and they were 
lacerated on both sides of the snout. Four were found 
dead. We released the four live individuals but they 
did not appear to be in good condition upon release; 


FiGuRE 1. Deer Mouse with its snout caught in the wire 
mesh of an Ugglan live-trap. The tin weather shield 
was removed for this photograph. 


we did not recapture these four individuals and sus- 
pect from their injury and constitution that they may 
have died sometime after release. All of the individu- 
als that were encountered with their snouts stuck in 
the wire mesh were found underneath the elevated trap 
door (Figure |), where the animals could see outside 
of the trap. We surmise that trapped Deer Mice were 
attempting to gnaw through the exposed wire mesh to 
escape and became caught. No other Deer Mice were 
found dead in the traps other than those with their 
snouts caught. 

During our sampling, we captured 443 individuals, 
representing five species (Deer Mice and 4 species of 
arvicoline rodents), a total of 888 times. None of the 
voles captured (Northern Red-backed Vole, Clethri- 
onomys rutilus; Long-tailed Vole, Microtus longi- 


445 


446 


caudus; Meadow Vole, M. pennslyvanicus; Taiga Vole, 
M. xanthognathus) captured were found with their 
snouts stuck in the wire mesh. Although some voles 
(12.9%, primarily juvenile C. rutilus) died in the traps 
from exposure, predation, or stress-related capture 
myopathy, we do not attribute this to trap type or tech- 
nique. We suggest that voles did not become entrapped 
like some Deer Mice because of the difference in their 
facial morphology; arvicoline rodents tend to have 
shorter and broader snouts than mice, thus their snout 
likely would not fit through the wire mesh of our live 
traps. 

The percentage of Deer Mice killed in our Ugglan 
traps is higher than reported in other studies of Per- 
omyscus. For example, Whittaker et al. (1998) report- 
ed that <1% (2 of 655) of Peromyscus (White-footed 
Mouse, P. leucopus and Cotton Mouse, P. gossypi- 
nus) captured had died in box-style (Sherman) traps. 
In Australia, Jacob et al. (2002) reported mortality rates 
of House Mice (Mus domesticus) in Ugglan traps as 
12%, wheras mortality in Longworth traps was only 
1%. They attributed mortality of house mice in Ugglan 
traps to exposure, however, and not to becoming caught 
in the wire mesh. 

This report of an unusual cause of trap-related mor- 
tality of Deer Mice in wire mesh traps is not intended 
to be a critique of wire mesh traps (which some believe 
to be a superior trap for some species of small mam- 
mals; e.g., O'Farrell et al. 1994). Rather, we wish to 
alert biologists intending to use similar traps of a 
potential problem. 

A smaller mesh size would likely prevent such mor- 
talities, but may be prohibitive to manufacture and 
would not address the use of those traps already in 
field use. We suggest that biologists using Ugglan traps 
can reduce the incidents of this type of trap mortality 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


by slightly modifying the traps. A small piece of card- 
board or tin can be placed to cover the wire mesh under 
the trap door without affecting it’s operation. This would 
likely reduce trap-related mortalities of mice and may 
also keep bait from sliding outside of the capture area. 
We note that some Ugglan traps came supplied from 
the manufacturer with such a tin shield, while others 
did not. 


Acknowledgments 

We thank C. Gagnon, V. Loewen, and J. Staniforth 
for field assistance. J. Adamczeski, V. Loewen, D. 
Nagorsen, M. O’ Donoghue, B. Slough and an anony- 
mous reviewer kindly suggested improvements to this 
note. Funding for the study that yielded these obser- 
vations was provided by the Yukon Department of 
Environment. 


Literature Cited 

Jacob, J., H. Ylonen, and C. G. Hodkinson. 2002. Trapping 
efficiency of Ugglan traps and Longworth traps for house 
mice in south-eastern Australia. Wildlife Research 29: 101- 
103. 

Jung, T. S., I. D. Thompson, M. B. C. Hickey, and R. D. 
Titman. 2002. Apparent capture myopathy in hoary bats, 
Lasiurus cinereus: a cautionary note. Canadian Field-Nat- 
uralist 116: 136-137. 

O’Farrell, M. J., W. A. Clark, F. H. Emmerson, S. M. 
Juarez, F. R. Kay, T. M. O’Farrell, and T. Y. Goodlett. 
1994. Use of a mesh live trap for small mammals: are 
results from Sherman live traps deceptive? Journal of 
Mammalogy 75: 692-699. 

Whittaker, J. C., G. A. Feldhamer, and E. C. Charles. 1998. 
Captures of mice, Peromyscus, in two sizes of Sherman 
live traps. Canadian Field-Naturalist 112: 527-529. 


Received 13 November 2003 
Accepted 15 February 2005 


Longevity and Productivity of Three Wolves, Canis lupus, in the Wild 


Jim Horyan,! DIANE K. Boyp,? Curtis M. Mack,! and DANIEL H. PLETSCHER? 


' Nez Perce Tribe Wolf Recovery Program, P.O. Box 1922, McCall, Idaho 83638 USA 
? Wildlife Biology Program, School of Forestry, University of Montana, Missoula, Montana 59828 USA 


Holyan, Jim, Diane K. Boyd, Curtis M. Mack, and Daniel H. Pletscher. 2005. Longevity and productivity of three Wolves, 
Canis lupus, in the wild. Canadian Field-Naturalist 119(3): 447-448. 


We document longevity and productivity of three Wolves (Canis lupus) in the wild in Montana and Idaho, USA. Two male 
Wolves each attained ages of at least 13 years, while a female was 12.2 years old. All three Wolves in our study were older 
at the time of their last known reproductive events than others reported in the literature. 


Key Words: Wolf, Canis lupus, longevity, reproduction, Montana, Idaho. 


Wolves (Canis lupus) are a relatively long-lived 
species, with individuals in captivity commonly reach- 
ing 9+ years, and occasionally living up to 17 years. 
(Young and Goldman 1944; Goodwin and Ballard 
1985; Okarma and Koteja 1987; Mech 1988; Landon 
et al. 1998; Mech and Boitani 2003). Mech (1988) 


stated that wild Wolves rarely exceed 13 years of age, 
and Peterson (1997:13), referring to the protected Wolf 
population in Isle Royale National Park, stated that 
maximum lifespan could reach “10-12 or even 14 
years,” though his statement was not supported with 
documentation. It is reasonable to assume that Wolves 


2005 


living in captivity would outlive wild Wolves facing 
more rigorous environments. 

Here, we report on three wild Wolves, 9013, 8756, 
and B-2, that each attained an estimated minimum age 
of 12.2 years. These animals were all radio-collared 
and inhabited Idaho and Montana, USA, from 1987 to 
2004. Data for these Wolves were collected using stan- 
dard radio-telemetry techniques in the Northern Rocky 
Mountain recovery areas of the USA (U.S. Fish and 
Wildlife Service et al. 2004). 

Ages of Wolves can be accurately determined for 
individuals < 2.5 years old based on tooth eruption and 
wear patterns (Gipson et al. 2000), body size, devel- 
opment and appearance of external reproductive organs, 
and previous known pack history. Using these crite- 
ria, and assuming a mid-April birth date (Boyd et al. 
1993; Thurston 2002), we were able to determine the 
ages for male Wolf 9013 and female Wolf 8756 at the 
times of their captures. We were only able to estimate 
male Wolf B-2s age because he was greater than 2.5 
years old at the time of his capture. 

Male Wolf 9013 was initially captured and radio- 
collared in September 1990 as a member of the South 
Camas pack in Glacier National Park, Montana, USA. 
We determined he was 2.4 years old at the time of his 
capture. Wolf 9013 dispersed from his natal pack in 
1991 to the Kelly Creek drainage of northern Idaho 
where he remained a lone wolf. In 1995, 15 Wolves 
were translocated into central Idaho as part of a fed- 
eral strategy to recover endangered Wolf populations 
in the Northern Rocky Mountains of the USA (U.S. 
Fish and Wildlife Service 1994*). By March 1996, 
Wolf 9013 paired had with female Wolf B-15, one of 
the Wolves translocated in 1995. They produced a lit- 
ter of five pups in 1997 and formed the Kelly Creek 
pack when Wolf 9013 was 8.8 years old. This breed- 
ing pair produced subsequent litters in 1998-2000. 
Wolf 9013 was 11.8 years old when he sired his final 
litter in 2000. 

Wolf 9013 may have been displaced as the alpha 
male by late 2000, as he was frequently located apart 
from other radio-collared members of his pack. The 
Kelly Creek pack failed to reproduce in 2001. Wolf 9013 
was 13.2 years of age when he died from unknown 
causes in June 2001. An incisor was collected for ag- 
ing (Goodwin and Ballard 1985). Cementum annuli 
analysis (Matson’s Laboratory, LLC) indicated an age 
of 8-9 years for Wolf 9013, which underestimated the 
known age by a minimum of 4.2 years, the error prob- 
ably caused by extensive cementum resorption (G. 
Matson, personal communication). 

Female Wolf 8756 was captured in Glacier Nation- 
al Park, Montana, USA, as a 5-month-old pup in Octo- 
ber 1987. She was monitored weekly with radio-teleme- 
try from 1987-1999. She became the breeding female 
of the South Camas Pack in Glacier National Park in 
1990 at the age of 2.8 years. She apparently produced 
a litter of pups every year and possibly produced her 


NOTES 


447 


last litter at 12 years of age. She was 12.2 years of age 
at the time of her death in June 1999 (Gipson et. al. 
2002). 

Male Wolf B-2 was estimated to be 4.8 years old 
when translocated to central Idaho during the federal 
recovery effort in 1995. Wolf B-2 paired with a dis- 
persing female Wolf and established the Wildhorse 
pack, where he fathered litters of pups in 2000 and 
2001, at the approximate ages of 9.8 and 10.8 years, 
respectively. He also sired a litter of four pups in 2003 
at approximately 12.8 years of age, after he established 
the Castle Peak pack. He died at the estimated age of 
13.8 years in February 2004. A lower first premolar 
was collected for aging (Goodwin and Ballard 1985). 
Cementum annuli analysis (Matson’s Laboratory, LLC) 
indicated an age of 11 years for Wolf B-2. 

The two male Wolves reported here lived longer 
than other longevous wild male Wolves reported in 
the literature; 9013 was 13.2 years old and B-2 was 
estimated at 13.8 years old. Mech (1988, 1997) reported 
minimum ages of 11.6 and 11.0 years for the two 
oldest males he studied. Female Wolf 8756 outlived all 
but one wild female Wolf reported by Mech (1988). 

Both males 9013 and B-2 sired litters at older ages 
than others reported in the literature. Mech (1988) noted 
the oldest male Wolf in Minnesota that sired pups was 
10.8 years of age; Wolf 9013 was 11.8 years of age in 
2000 when he was still believed to be the breeding male 
of his pack. Wolf B-2 was an estimated 12.8 years old 
when he was known to be the breeding male of his 
pack. 

Kreeger (2003:193) stated “maximum breeding age 
for female wolves is not known.” Female Wolf 8756 
exceeded the maximum breeding age previously report- 
ed when she produced pups at 11 years of age, and she 
may have reproduced at 12 years old. 

The value of known-age animals is significant in 
describing longevity. Estimating age with cementum 
annuli analysis may not be accurate for Wolves (Good- 
win and Ballard 1985; Landon et al. 1998). Gipson et 
al. (2000) concluded that cementum annuli analysis 
underestimated actual age by 1-3 years for Wolves >3- 
14 years of age. Our longevity data for two Wolves of 
known age and a third with a good age estimation, and 
their reproductive activity at advanced ages, are note- 
worthy. 


Acknowledgments 

The University of Montana, the U.S. Fish and Wild- 
life Service, the National Park Service, and the Nez 
Perce Tribe supported this study. We thank I. Babcock, 
M. Carter, A. Sondenaa, and two anonymous review- 
ers for comments that improved this manuscript. 


Documents Cited (marked * in text) 

U.S. Fish and Wildlife Service. 1994. The reintroduction 
of gray wolves to Yellowstone National Park and central 
Idaho: Final environmental impact statement. U.S. Fish 
and Wildlife Service, Helena, Montana. 562 pages. 


448 


Literature Cited 

Boyd, D. K., R. R. Ream, D. H. Pletscher, and M. W. Fair- 
child. 1993. Variation in denning and parturition dates of 
a wild gray wolf, Canis lupus, in the Rocky Mountains. 
Canadian Field-Naturalist 107: 359-360. 

Gipson, P. S., W. B. Ballard, R. M. Nowak, and L. D. Mech. 
2000. Accuracy and precision of estimating age of gray 
wolves by tooth wear. Journal of Wildlife Management 
64: 752-758. 

Gipson, P. S., E. E. Bangs, T. N. Bailey, D. K. Boyd, H. D. 
Cluff, D. W. Smith, and M. D. Jimenez. 2002. Color 
patterns among wolves in western North America. Wild- 
life Society Bulletin 30: 821-830. 

Goodwin, E. A., and W. B. Ballard. 1985. Use of tooth ce- 
mentum for age determination of gray wolves. Journal of 
Wildlife Management 49: 313-316. 

Kreeger, T. J. 2003. The internal wolf: physiology, pathology, 
and pharmacology. Pages 192-217 in Wolves: Behavior, 
ecology, and conservation. Edited by L. D. Mech and L. 
Boitani. University of Chicago Press, Chicago, Illinois. 
448 pages. 

Landon, D. B., C. A. Waite, R. O. Peterson, and L. D. Mech. 
1998. Evaluation of age determination techniques for gray 
wolves. Journal of Wildlife Management 62: 674-682. 

Mech, L. D. 1988. Longevity in wild wolves. Journal of Mam- 
malogy 69: 197-198. 

Mech, L. D. 1997. An example of endurance in an old wolf, 
Canis lupus. Canadian Field-Naturalist 111: 654-655. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Mech, L. D., and L. Boitani. Editors. 2003. Wolves: Behavior, 
ecology, and conservation. University of Chicago Press, 
Chicago, Illinois. 448 pages. 

Okarma, H., and P. Koteja. 1987. Basal metabolic rate in the 
gray wolf in Poland. Journal of Wildlife Management 51: 
800-801. 

Peterson, R. O. 1997. The wolves and moose of Isle Royale. 
Pages 6-14 in World Wide Fund for Nature’s Vargsympo- 
sium 1997. Edited by L. Nyman and M. Ericson Varlds- 
naturfonden WWE 3:98. 

Thurston, L. M. 2002. Homesite attendance as a measure 
of alloparental and parental care by gray wolves (Canis 
lupus) in northern Yellowstone National Park. MA thesis, 
Texas A & M University, College Station, Texas. 175 pages. 

U.S. Fish and Wildlife Service, Nez Perce Tribe, National 
Park Service, Montana Fish, Wildlife and Parks, Idaho 
Fish and Game, and U.S.D.A. Wildlife Services. 2004. 
Rocky Mountain Wolf Recovery 2003 Annual Report. 
Edited by D. Boyd. USFWS Ecological Services, Helena, 
Montana. 72 pages. 

Young, S. P., and E. A. Goldman. 1944. The wolves of 
North America. Dover Publishing, New York, New York. 
385 pages. 


Received 10 May 2004 
Accepted 6 June 2005 


2005 NOTES 449 


Observations of Autumn Courtship and Breeding in Brown Bears, 
Ursus arctos, from Coastal British Columbia 


OwEN T. NEVIN! and BARRIE K. GILBERT? 


'Centre for Animal Conservation, University of Central Lancashire, Penrith, Cumbria, CA1 1 OAH, United Kingdom 
"Department of Forest, Range and Wildlife Sciences, Utah State University, Logan, Utah 84322 USA 


Nevin, Owen T., and Barrie K. Gilbert. 2005. Observations of autumn courtship and breeding in Brown Bears, Ursus arctos, 
from coastal British Columbia. Canadian Field-Naturalist 119(3): 449-450. 


Over a period of four years, autumn courtship behaviour in Brown Bears (Ursus arctos) was observed on three separate 
occasions, with copulation observed once and two litters of cubs potentially resulting from autumn breeding. These are the 


first recorded observations of these behaviours. 


Key Words: Brown Bear, Ursus arctos, reproduction, autumn, courtship, mating, British Columbia. 


Brown Bears (Ursus arctos) mate in the spring from 
May to July. Once the fertilized eggs have developed to 
the blastocyst stage, further development is delayed 
until just before denning, typically November. At this 
time the blastocysts implant and a short pregnancy 
ensues with one to four cubs born during winter sleep, 
usually in February. This delayed implantation leads to 
an apparent gestation period of 6.5 — 8.5 months, 
although embryonic development occurs only in the last 
6 — 8 weeks (Craighead et al. 1969; Hensel et al. 1969; 
Garshelis 2001). 

Between August 1999 and May 2003, Brown Bears 
in the Glendale Cove area of British Columbia (50°41'N 
125°44'W) were individually identified and observed as 
part of an ongoing study (Nevin 2003). Photo-identifi- 
cation techniques allowed individual bears to be distin- 
guished. Coat coloration and scar patterns were record- 
ed with sketches and descriptions on data sheets, 
supplemented by a catalogue of reference photo- 
graphs. Each bear was given a unique numeric code. 
Sex was determined by urination pattern, direct 
observation of genitals or the presence of cubs 
(Nevin 2003; Nevin and Gilbert 2005a). 

Autumn courtship was first observed in mid-Septem- 
ber 1999; two large adult males (M003 and M004) 
were observed tracking a lone female (F005). While 
this close following and olfactory investigation of 
urine is typical of the breeding season, it was only in 
light of subsequent observations that this was recog- 
nized as out-of-season courtship behaviour. 

Two weeks later, at the beginning of October 1999, 
one of these male bears (M003) was observed courting 
another lone female (F102). During the 40 minutes that 
the interactions of these bears were recorded, tracking 
was again observed; in addition, there was play fighting, 
genital sniffing and mounting. 

In the same week one of the female bears (F009) lost 
two yearling cubs. The cubs were last seen on 3 Octo- 
ber 1999 after which FO09 was seen alone. F009 is a 
highly identifiable bear with distinctive scarring and 
a muzzle that twists to the right. After the loss of her 
cubs in 1999 she was not seen in 2000 but returned to 


the Glendale spawning channel in early September 
2001. At this time she was accompanied by three year- 
ling cubs; these cubs would have been born during 
the 1999-2000 hibernation. For this to happen, FO0O9 
must either have mated in the spring of 1999 while still 
with her yearling cubs or have mated in the autumn of 
1999 after the loss of her cubs. Neither explanation 
would be considered typical behaviour. 

In mid September 2001 M004 was again seen track- 
ing FOOS; this is the same pairing seen in September 
1999 and FOOS still had no cubs. On this occasion 
they approached another female (F103) and her two- 
year-old cubs. F103 charged FOOS and after a brief 
agonistic interaction F005, the larger of the females, 
backed away approximately 10 m and sat down. At this 
point F103 was approached by M004; after another 
brief agonistic interaction, F103 moved away from her 
cubs and offered no resistance to mounting by M004. 
Copulation began immediately and continued for at 
least 10 minutes at which time continued observation 
became impossible. This is the first recorded observa- 
tion of autumn copulation in Brown Bears. It should 
be noted that F103 had been involved in courtship 
behaviour in the spring (e.g. 15 May 2001 with M301); 
she had separated from her cubs in late May and was 
seen mating with M301 in May and June. F103 and 
cubs had reunited by 16 September 2001. F103 was 
seen in May 2002 with two young-of-year cubs but it 
is not known whether these are from spring or autumn 
mating in 2001. 

With only four observations over a period of four 
years during which 341 days of systematic behavioural 
observations were conducted for ongoing research proj- 
ects (Nevin 2003; Nevin and Gilbert 200Sa, b) on the 
approximately 40 individual bears using the area, we 
are Clearly discussing rare events; during this period 
more than 30 breeding events were observed during the 
spring breeding season. It should be noted that although 
all the observations of autumn courtship and breed- 
ing involve one of two male bears (M003 and M004) 
these were two of the five largest males in the area. 

Very little is known about the reproductive biology 
of any bear species and new insight into Brown Bear 


450 


breeding may have impacts on the conservation and 
management of other bear species, many of which are 
threatened or endangered (Spady 2002). The potential 
of successful autumn breeding will also add a new 
dimension to the ongoing debate on infanticide in 
Brown Bears (Wielgus et al. 2000, 2001; Swenson et 
al. 2001a, b; Dahle and Swenson 2003; Miller et al. 
2003; Ben-David et al. 2004; Nevin and Gilbert 2005a, 
b). With the possibility of same-season breeding oppor- 
tunities associated with cub-killing events throughout 
the non-denning period, the selective advantage asso- 
ciated with this behaviour would be much higher 
than it has been assumed to be under the prevailing 
view that breeding is restricted to the spring period. 


Acknowledgments 

This research was funded through ongoing support 
from Knight Inlet Lodge and the Glendale Grizzly 
Trust. Special thanks go to Barbara Bull, Dean Wyatt 
and the staff of Knight Inlet Lodge. 


Literature Cited 

Ben-David, M., K. Titus, and L. R. Beier. 2004. Consump- 
tion of salmon by Alaskan brown bears: A trade-off between 
nutritional requirements and the risk of infanticide? Oeco- 
logia 138: 465-474. 

Craighead, J. J.. M. G. Hornocker, and F. C. Craighead, 
Jr. 1969. Reproductive biology of young female grizzly 
bears. Journal of Reproduction and Fertility Supplement 
6: 447-475. 

Dahle, B, and J. E. Swenson. 2003. Seasonal range size in 
relation to reproductive strategies in brown bears Ursus 
arctos. Journal of Animal Ecology 72: 660-667. 

Garshelis, D. L. 2001. Bear family. Pages 70-85 in The new 
encyclopedia of mammals. Edited by D. Macdonald. Ox- 
ford University Press, Oxford. 


THE CANADIAN FIELD-NATURALIST 


Vol. P89 


Hensel, R. J., W. A. Troyer, and A. W. Erickson. 1969. 
Reproduction in the female brown bear. Journal of Wild- 
life Management 33: 357-365. 

Miller, S D., R. A. Sellers, and J. A. Keay. 2003. Effects of 
hunting on brown bear cub survival and litter size in Alas- 
ka. Ursus 14: 130-152. 

Nevin, O. T. 2003. The influence of prey abundance and 
risk-sensitive behavioral change on individual access to 
high-energy food (salmon): impacts on the density and 
viability of bear populations. Ph.D. dissertation, Utah State 
University, Logan, Utah. 

Nevin, O. T., and B. K. Gilbert. 2005a. Perceived risk, dis- 
placement and refuging in brown bears: positive impacts 
of ecotourism? Biological Conservation 121: 611-622. 

Nevin, O. T., and B. K. Gilbert. 2005b. Measuring the cost 
of risk avoidance in brown bears: Further evidence of posi- 
tive impacts of ecotourism. Biological Conservation 123: 
453-460. 

Spady, T. 2002. Joint ursid reproduction study. International 
Bear News 11(3): 14. 

Swenson, J. E., B., Dahle, and F. Sandegren. 200 1a. Intra- 
specific predation in Scandinavian brown bears older than 
cubs-of-the-year. Ursus 12: 81-92. 

Swenson, J. E., F. Sandegren, S. Brunberg, and P. Seger- 
strom. 2001b. Factors associated with loss of brown bear 
cubs in Sweden. Ursus 12: 69-80. 

Wielgus, R. B., and F. L. Bunnell. 2000. Possible negative 
effects of adult male mortality on female grizzly bear re- 
production. Biological Conservation 93: 145-154. 

Wielgus, R. B., F. Sarrazin, R. Ferriere, and J. Clobert. 
2001. Estimating effects of adult male mortality on grizzly 
bear population growth and persistence using matrix models. 
Biological Conservation 98: 293-303. 


Received 9 August 2004 
Accepted 22 August 2005 


2005 


NOTES 


45] 


Long-distance Movement of a Dispersing Deer Mouse, Peromyscus 


maniculatus, 1n the Boreal Forest 


Thomas S. Jung!, Kieran S. O’Donovan!, and Todd Powell? 


' Yukon Department of Environment, P.O. Box 2703, Whitehorse, Yukon Y1A 2C6 Canada 
? Yukon Department of Environment, P.O. Box 194, Watson Lake, Yukon YOA 1CO Canada 


Jung, Thomas S., Kieran S. O’Donovan, and Todd Powell. 2005. Long-distance movement of a dispersing Deer Mouse, 
Peromyscus maniculatus, in the boreal forest. Canadian Field-Naturalist 119(3): 451-452. 


We report an apparent long-distance, non-homing movement of 3044 + 60 m made by a dispersing subadult male Deer 
Mouse, Peromyscus maniculatus, in southeastern Yukon. Our observation is nearly twice the maximum distance previously 
recorded for non-homing Deer Mice, and apparently the longest dispersal movement recorded for this species. 


Key Words: Deer Mouse, Peromyscus maniculatus, dispersal, movements, Yukon. 


Despite the significance of dispersal in the spatial 
ecology and population dynamics of small mammals, 
the process is little understood (Fairbairn 1978; Bow- 
man et al. 2001). Deer Mice (Peromyscus maniculatus) 
can move relatively long distances (e.g. >1000 m; 
Teferi and Millar 1993; Topping et al. 1997; Bowman 
et al. 1999, 2001; Rehmeier et al. 2004), yet reports of 
such movements are rare, likely due to a low probability 
of detection (Rehmeier et al. 2004). Here, we report an 
apparent record long-distance movement by a dispers- 
ing Deer Mouse in southeastern Yukon. 

As a part of studies on small mammals in the boreal 
forest near Watson Lake, Yukon (60.06°N, 128.70°W), 
we used Ugglan live-traps (Model 3 Lemming Special, 
Granhab, Marieholm, Sweden) and Longworth live- 
traps (Longworth Scientific Instruments Co., Abing- 
don, United Kingdom) to live-capture small mam- 
mals during summer and fall 2004. We live-trapped 
rodents at two study areas situated about 3 km apart. 
One trapping area (Area A) was located about | km 
N of Watson Lake, while the other (Area B) was 
about 3 km NE of Watson Lake. Both study areas were 
in boreal mixedwood forest. Co-dominant tree species 
included White Spruce (Picea glauca), Lodgepole Pine 
(Pinus contorta), Trembling Aspen (Populus tremu- 
loides) and White Birch (Betula papyrifera). Limited 
logging occurred in Area B 8-12 years prior to our 
trapping. 

At Area A, we established 6 grids of 25 trapping 
stations (5 x 5 layout; 10 m between traps). We placed 
one Ugglan and one Longworth trap at each trapping 
station. Traps were operational for eight days with no 
pre-baiting. At Area B, we ran 14 grids of 48-49 trap- 
ping stations (6 x 8 or 7 x 7 layout; 12 m between 
traps). One Ugglan trap was placed at each trapping 
station. Traps on these grids were pre-baited for two 
days and operational for three days. At both study 
areas, traps were 12 m apart, baited with rolled oats, 
peanut butter and a slice of carrot, and cotton bedding 
was provided. Each morning, captured animals were 
identified, weighed, sexed, aged, ear-tagged with a 
Number | Monel tag (National Band and Tag Com- 


pany, Newport, Kansas), and then released where they 
were caught. 

On 11 August 2004, we captured a subadult male 
Deer Mouse (eartag # 1027) in a Longworth trap at 
Area A. On 26 August 2004, 15 days after being orig- 
inally captured at Area A, the mouse was recaptured in 
an Ugglan trap at Area B. Geographic co-ordinates (+ 
30 m) of the original capture location and the last cap- 
ture location were obtained with a global positioning 
system (GPS; Garmin International Inc., Olathe, 
Kansas). We used a geographic information system 
(GIS; ArcView GIS 3.2, ESRI, Redlands, California) to 
calculate the distance moved by the Deer Mouse. GIS 
analysis provided a straight-line distance of 3044 
+ 60 m between the two capture locations. Travel by the 
Deer Mouse was most likely through continuous boreal 
forest. The straight-line movement, or substantial varia- 
tions thereof, indicate that the movement between cap- 
ture locations was neither hindered nor facilitated by 
apparent barriers (e.g., rivers) or travel corridors (e.g., 
powerlines or roads), respectively. 

No other investigators had trapped and ear-tagged 
small mammals in the area in the past five years, so it is 
unlikely that the Deer Mouse was from another study. 
Furthermore, our ear tags were prefaced with the code 
“YTG” before the numbering, which is distinctive in 
the field. Similarly, we are confident that the tag was not 
confused with another Deer Mouse ear-tagged on the 
grid where it was last recaptured, since none of the 
other five Deer Mice captured at this grid had tag num- 
bers that were easily confused with # 1027. We did not 
simultaneously trap at both study areas, and different 
field crews were doing the work at the two study areas, 
so we consider it unlikely that the mouse traveled in a 
vehicle or trap with a field crew from Area A to Area B. 
Given rejection of these potential alternatives, we con- 
clude that the Deer Mouse did, in all likelihood, move 
a minimum distance of 3044 + 60 m. 

Bowman et al. (1999) recorded a long-distance dis- 
persal movement of 1768 m by a subadult male Deer 
Mouse in forests in New Brunswick. In Kansas, 
Rehmeier et al. (2004) reported several Deer Mice mov- 


452 


ing > 1000 m, with a maximum recorded movement of 
1320 m. During homing experiments, several studies 
have reported human-induced movements of Deer Mice 
>1500 m (Murie 1963; Furrer 1973; Teferi and Millar 
1993). In Alberta, Teferi and Millar (1993) reported one 
Deer Mouse had moved a straight-line distance of 
1980 m during a homing experiment. Maier (2002) 
reported a female White-footed Mouse (Peromyscus 
leucopus) naturally moving a minimum of 14730 m, 
and another female White-footed Mouse moving 
6840 m, between successive capture locations in Massa- 
chusetts. Therefore, our observation of a minimum 
movement of >3000 m for a Deer Mouse is not incon- 
ceivable. To the best of our knowledge, our observation 
establishes the longest recorded movement of a Deer 
Mouse. 

Given the time of year (early fall) and that the indi- 
vidual was a subadult male, we suggest that the ob- 
served movement represents dispersal from the natal 
range (Fairbairn 1978). The mechanisms behind dis- 
persal in Peromyscus are unknown. Rodent populations 
may vary widely from year to year (e.g., Gilbert and 
Krebs 1991), and Bowman et al. (1999) and Maier 
(2002) had suggested that long-distance dispersal in 
Peromyscus in their study areas may have been influ- 
enced by high population densities. In our study, how- 
ever, Deer Mouse abundance was 3.4 times lower in 
2004 than in 2003; 2.01 individuals per 100 trap nights 
(TN) vs. 6.77 individuals per 100 TN, respectively (Jung 
et al. unpublished data). Therefore, there should have 
been much nearby, suitable, and unoccupied habitat 
for the Deer Mouse to establish a territory. Rehmeier 
et al. (2004), using multiple observations over nine 
years, also noted an inverse relationship between popu- 
lation density and propensity of Deer Mice to move 
long distances. They suggested that increased num- 
bers of social fences at higher densities may result in 
decreased incidences of long distance movements by 
Deer Mice. Maier (2002) suggested that a lack of tra- 
ditionally available food resources may also lead to 
notably long-distance dispersal. We have no data on 
food availability for murine rodents in our study area. 
Regardless of the mechanism or trigger, our observa- 
tion substantiates the contention by Bowman et al. 


~ THE CANADIAN FIELD-NATURALIST 


Vol. 119 


(1999), Rehmeier et al. (2004), and Maier (2002) that 
small mammals may be more vagile than previously 
thought. 


Acknowledgments 

We thank K. Russell, J. Colbert, and R. Brodhagen 
for field assistance. M. O’Donoghue, B. Slough, D. 
Nagorsen and two anonymous reviewers kindly re- 
viewed an earlier draft of this note. We thank the Yukon 
Department of Environment for its continued support 
of wildlife research. 


Literature Cited 

Bowman, J. C., M. Edwards, L. S. Sheppard, and G. J. 
Forbes. 1999. Record distance for a non-homing move- 
ment by a Deer Mouse, Peromyscus maniculatus. Cana- 
dian Field-Naturalist 113: 292-293. 

Bowman, J. C., G. J. Forbes, and T. G. Dilworth. 2001. 
Distances moved by small woodland rodents within large 
trapping grids. Canadian Field-Naturalist 115: 64-67. 

Fairbairn, D. J. 1978. Dispersal of Deer Mice, Peromyscus 
maniculatus: proximal causes and effects on fitness. Oecol- 
ogia 32: 171-193. 

Furrer, R. K. 1973. Homing of Peromyscus maniculatus in 
the Channelled Scablands of east-central Washington. Jour- 
nal of Mammalogy 54: 466-483. 

Gilbert, B. S., and C. J. Krebs. 1991. Population dynamics 
of Clethrionomys and Peromyscus in southwestern Yukon, 
1973-1989. Holarctic Ecology 14: 250-259. 

Maier, T. J. 2002. Long-distance movements by female white- 
footed mice, Peromyscus leucopus, in extensive mixed- 
wood forest. Canadian Field-Naturalist 116: 108-111. 

Murie, M. 1963. Homing and orientation of Deer Mice. 
Journal of Mammalogy 44: 338-349. 

Rehmeier, R. L., G. A. Kaufman, and D. W. Kaufman. 
2004. Long-distance movements of the Deer Mouse in 
tallgrass prairie. Journal of Mammalogy 85: 562-568. 

Teferi, T., and J. S. Millar. 1993. Long distance homing by 
the Deer Mouse, Peromyscus maniculatus. Canadian Field- 
Naturalist 107: 109-111 

Topping, M. G., J. S. Millar, and B. E. Woolfenden. 1997. 
Unsuccessful colonization of a naturally depopulated area 
by the Deer Mouse, Peromyscus maniculatus. Canadian 
Field-Naturalist 111: 466-469. 


Received 17 September 2004 
Accepted 8 September 2005 


2005 NOTES 453 


Movements of Two Rabid Raccoons, Procyon lotor, in Eastern Ontario 


Rick RosaTtTe!3, MIKE ALLAN!, ROB WARREN!, PETER NEAVE!, TopD BABIN!, LUKE BUCHANAN!, 
DENNIS DONOVAN!, KIRK SoBEY!, CHRIS DAviEs!, FRANCES MULDOON’, and ALEX WANDELER? 


‘Ontario Ministry of Natural Resources, Wildlife Research and Development Section, Trent University, Science Complex, 
P.O. Box 4840, Peterborough, Ontario K9J 8N8 Canada 

*Canadian Food Inspection Agency, Ottawa Laboratory Fallowfield, P.O. Box 11300, Station H, Nepean, Ontario K2H 8P9 
Canada 

3Corresponding author: e-mail: rick.rosatte@mnr.gov.on.ca); Tel. (705) 755-2280, Fax (705) 755-2276 


Rosatte, Rick, Mike Allan, Rob Warren, Peter Neave, Todd Babin, Luke Buchanan, Dennis Donovan, Kirk Sobey, Chris Davies, 
Frances Muldoon and Alex Wandeler. 2005. Movements of two rabid Raccoons, Procyon lotor, in Eastern Ontario. 
Canadian Field-Naturalist 119(3): 453-454. 


An adult female Raccoon Procyon lotor was captured about 3 km north of Mallorytown, Ontario, on 27 August 2004, as part 
of a government rabies control program. The animal was vaccinated against rabies, ear-tagged and released, and recaptured the 
next day 1.7 km south of the initial capture location. Upon recapture, the Raccoon had porcupine quills in its facial area and 
seemed agitated and was submitted for rabies testing. It was confirmed as rabies positive on 31 August 2004, by the Canadian 
Food Inspection Agency. Similarly, a juvenile male raccoon was captured, ear-tagged, vaccinated, and released near Junetown, 
Ontario (about 4 km NW of the other rabid Raccoon) on 5 September 2004. It was found dying in a residential window well on 


22 September 2004, 700 meters from the original capture location. It was diagnosed as rabid on 23 September 2004. 


Key Words: Raccoon, Procyon lotor, rabies, movement, disease transmission, Ontario. 


The Raccoon, Procyon lotor, variant of rabies was first 
reported in Ontario, Canada, in July 1999 and 131 cases 
have been reported to September 2005 (Wandeler and 
Salsberg 1999; Rosatte et al. 2001). In response to 
these cases, the Ontario Ministry of Natural Resources 
(OMNR) implemented three different tactics to con- 
trol the disease: Trap-Vaccinate-Release (TVR), Point 
Infection Control (PIC), and Oral Rabies Vaccination 
with baits (ORV) (Rosatte et al. 2001). The use of TVR 
involves live-capture (with humane cage traps) and 
vaccination with an injection of Imrab rabies vaccine 
(Merial, Athens, Georgia). Release of target animals 
such as Raccoons allows researchers to gain informa- 
tion on movements if the animals ear-tagged for iden- 
tification during processing are recaptured. In addition, 
data regarding the movement of rabid Raccoons is 
scarce in the literature. 

On 27 August 2004, an adult female Raccoon was 
captured (Tomahawk live-trap, Tomahawk, Wisconsin) 
(about 3 km north of Mallorytown, Ontario), as part of 
a TVR program in response to two cases of Raccoon 
strain rabies that had occurred near Mallorytown during 
mid-August 2004. This Raccoon appeared normal 
except it was vocalizing repeatedly with high-pitched 
whining sounds. A young female Raccoon was also 
captured at the same time about 2 meters from the adult 
female Raccoon and was probably one of her litter (as 
the adult female was lactating). Both Raccoons were 
ear-tagged, vaccinated against rabies (Imrab) and 
released at the point of capture. A Global Positioning 
System (GPS) reading (Magellan Trailblazer 300) was 
taken at the location where the animal was captured 
(430112E, 4927926N). The next day, 28 August 2004, 
the adult female Raccoon was recaptured at GPS 


430094E, 4926235N, 1.7 km south of the initial capture 
location. On this occasion, the Raccoon was very agi- 
tated — constantly moving back and forth in the live- 
trap, turning upside down repeatedly, vocalizing with a 
high-pitched whine, and had three Porcupine, Erethi- 
zon dorsatum, quills in its facial area (snout). This 
Raccoon was humanely euthanized (via an injection) 
and submitted to the Canadian Food Inspection Agency, 
Ottawa Laboratory Fallowfield (CFIA OLF) in Nepean, 
Ontario, for rabies testing. Another lactating adult fe- 
male Raccoon and two young female Raccoons were 
also captured at this time within 4 meters of the quilled 
Raccoon. These three Raccoons were ear-tagged, vac- 
cinated against rabies with Imrab and released. The 
quilled adult female Raccoon was confirmed at CFIA 
OLF as positive for rabies on 31 August 2004. On | Sep- 
tember 2004 the virus was determined at CFIA OLF to 
be the Raccoon variant of rabies. 

On 5 September 2004, a juvenile male Raccoon was 
captured, ear-tagged, vaccinated, and released near 
Junetown, Ontario (about 4.2 km NW of the other rabid 
Raccoon at GPS 426300E 4928200N ). The Raccoon 
appeared normal at the time. Several other Raccoons 
were also captured at this location. It was found dying 
in a residential window well on 22 September 2004, 
700 meters from the original capture location. This rac- 
coon was diagnosed as rabid on 23 September 2004. 

Data regarding the movement of rabid Raccoons is 
critical when designing rabies control tactics, especially 
when determining the width of vaccination zones to 
contain the disease (Rosatte et al. 2001). Even less is 
known about the contact rates of rabid Raccoons with 
susceptible animals. These rates will, in turn, affect the 
intensity and movement of a rabies outbreak (Totton et 


454 


al. 2002). Due to the close proximity of the rabid Rac- 
coon to the four other Raccoons that were captured, as 
well as the Porcupine quills in the rabid Raccoon (no 
rabid Porcupines have been reported in the area), there 
was great potential for this Raccoon to have transmitted 
rabies to multiple individuals, both intra-specifically as 
well inter-specifically, during its journey of at least 1.7 
km (assuming the Raccoon moved in a straight line 
which is unlikely). Analysis of the mark-recapture data 
from the area in which the rabid Raccoon was captured 
indicated that the average Raccoon density was 10 rac- 
coons/km’. Given this high Raccoon density, the poten- 
tial for rabies transmission to susceptible individuals in 
the area where the rabid Raccoon travelled was prob- 
ably very high. This information was subsequently used 
to modify the OMNR’s response to Raccoon rabies in 
eastern Ontario; the area where the rabid animal was 
captured was re-trapped and all abnormally acting Rac- 
coons and Striped Skunks (Mephitis mephitis) were 
humanely euthanized and submitted for rabies testing. 
All animals that had been vaccinated during previous 
years were re-vaccinated against rabies and released. 

Also of importance is the fact that both of these Rac- 
coons were vaccinated against rabies, though vaccina- 
tion did not protect them from developing clinical 
rabies. This in all likelihood was due to the fact that the 
first Raccoon already showed prodromal symptoms at 
the time of vaccination and the other Raccoon was incu- 
bating the disease. This emphasizes the value of Point 
Infection Control programs using population reduction 
around previous cases of rabies to remove clinical and 
incubating animals from the population — i.e., had 
population reduction been utilized instead of TVR both 
of these Raccoons would have been humanely eutha- 
nized and removed from the population negating any 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


potential for them to infect other animals. On the other 
hand, population reduction would also have killed a 
large number of healthy animals that were already vac- 
cinated against rabies from aerial baiting or previous 
TVR work. In this particular instance TVR provided 
valuable information, though euthanasia would have 
been the preferred disease control option. Obviously, 
both TVR and population reduction have their advan- 
tages when used for the control of wildlife rabies. 


Acknowledgments 

The Ontario rabies program is supported by the 
Provincial Rabies Advisory Committee, Dr. J. Carlson, 
Chair, and the Ontario Ministry of Natural Resources 
(OMNR), Wildlife Research and Development Section, 
Dr. C. Davies, Manager. Special acknowledgment goes 
to the staff of the OMNR, Rabies Research and Devel- 
opment Unit in Peterborough and the CFIA, OLF 
Rabies Laboratory, Nepean. This is OMNR Wildlife 
Research and Development Section contribution num- 
ber 04-05. 


Literature Cited 

Rosatte, R., D. Donovan, M. Allan, L. Howes, A. Silver, 
K. Bennett, C. MacInnes, C. Davies, A. Wandeler, and 
B. Radford. 2001. Emergency response to raccoon rabies 
introduction into Ontario. Journal of Wildlife Diseases 
372 265-279" 

Totton, S. C., R. R. Tinline, R. C. Rosatte, and L. L. 
Bigler. 2002. Contact rates of raccoons Procyon lotor at a 
communal feeding site in rural eastern Ontario. Journal 
of Wildlife Diseases 38: 313-319. 

Wandeler, A., and E. Salsberg. 1999. Raccoon rabies in 
eastern Ontario. Canadian Veterinary Journal 40: 731. 


Received 27 September 2004 
Accepted 6 September 2005 


2005 


NOTES 


455 


Common Loon, Gavia immer, Nest Attendance Patterns Recorded 


by Remote Video Camera 


WING GOODALE, LEE ATTIX, and DAVID EVERS 


BioDiversity Research Institute, 19 Flaggy Meadow Road, Gorham, Maine 04038 USA 


Goodale, Wing, Lee Attix, and David Evers. 2005. Common Loon, Gavia immer, nest attendance patterns recorded by 
remote video camera. Canadian Field-Naturalist 119(3): 455-456. 


We recorded the complete day/night nesting attendance of a Common Loon (Gavia immer) pair using a remote video camera. 
We found that the male and female share incubation duty equally, but that the female incubates primarily at night (95.8%) 


when the male defends the territory. 


Key Words: Common Loon, Gavia immer, incubation pattern, sex, day, night, remote camera, Maine. 


Researchers in the past have been limited to diurnal 
observations of Common Loon (Gavia immer) nesting 
behavior. Whereas many of those studies were con- 
ducted on unmarked loons (Sjolander and Agren 1972; 
Taylor 1974; McIntyre 1975), making the role of sex in 
incubation difficult to study, recent capture and mark- 
ing techniques by Evers (1994, 2001) now allow re- 
searchers to determine the sex of the incubating birds. 
Past studies of nest attendance with marked birds have 
shown possible relationships between time of day and 
sex of the incubating bird. Paruk (2000) found that 23% 
of the time there was a least one nest switch during the 
night, suggesting nest exchanges may be “optimized” 
for males to patrol the territory at night. Gostomski 
and Evers (1998) found that males incubated twice as 
often in the evening and the females twice as often in 
the morning, indicating the pair switches incubation 
duty after sunset. Evers (1994) found no significant sex 
difference in incubation effort during daylight hours. 
Our study explored parental incubation patterns by us- 
ing a remote video camera to record both day and night 
nest attendance of a marked Common Loon pair. 

In 2001 the Common Loon pair on Coleman Pond, 
Maine (44°53'25", 70°45'32") was color-banded and 
sexed (through morphometric measurements) using body 
mass as the primary criterion (Evers 2004*). On April 
2003 we set up a remote video camera equipped with 
a standard lens as well as a infrared lens and light for 
night vision (supplied by SeeMore Wildlife Systems) 
at a traditional nesting island in Coleman Pond, which 
is a single loon-territory lake. The camera, powered by 
solar panels, sent live images to a base computer and 
time-lapse VCR. At dusk the infrared light and lens 
would automatically turn on, allowing a recording of 
the entire incubation period. We then reviewed the 
time-lapse tapes and recorded (into a Microsoft Excel 
spreadsheet) the sex of the incubating bird (using the 
color bands) and the time of each nest exchange, as 
well as the amount of time the nest was left unattended. 
Events were not recorded after the eggs began pipping. 
Exact sunrise and sunset times for the lake were ob- 
tained from U.S. Navy records. From these data we 
calculated the precise time the male and female were 
on the nest during the day and night. 


TABLE 1. Total hours and minutes of nest attendance by 
time of day and sex (percentage is in parentheses). 


Empty Female Male Total 
Day  30:30(7.5) — 97:08 (23.8) 280:58 (68.8) 408:36 
Night, .<0:55'(0:5) 193:39:(95.8) TS3NS FT), go V207 
Total 31:25(5.1) 290:47 (47.6) 288:31 (47.2) 610:43 


Our results demonstrated that male and female loons 
shared incubation duty equally (Table 1). However, the 
female incubated the eggs primarily at night, whereas 
the male incubated primarily during the day. Gostom- 
ski and Evers (1998) interpreted that the male/female 
dusk and dawn nest attendance pattern indicated that 
the male defended the territory at night. That observa- 
tion confirmed findings by McIntyre (1988), who sug- 
gested that during incubation males patrol the territory 
at night. Our results support those findings, and qual- 
itative observations of the Coleman Pond loon pair 
indicated the male actively defended the territory at 
night from intruder loons with frequent yodel calls 
while the female was incubating. 


Acknowledgments 

We especially thank the MBNA Foundation, whose 
financial support made this project possible. In addition, 
Whitney and Tony Oppersdorff deserve special thanks 
for their generousity in providing a base station for 
the research equipment. Michelle Gharst also deserves 
special recognition for her review of the tapes. 


Document Cited (marked * in text) 

Evers, D. C. 2004. Status assessment and conservation plan for 
the Common Loon (Gavia immer) in North America: Final 
Draft. U.S. Fish and Wildlife Service, Hadley, Massachu- 
setts. 


Literature Cited 

Evers, D. C. 1994. Activity budget of marked Common Loon 
(Gavia immer) nesting population. Hydrobiologia 279/280: 
415-420. 

Evers, D. C. 2001. Capture and color-marking Common 
Loons: an evaluation of potential impacts. Pages 2-26 in 
Common Loon population studies: continental mercury 


456 


patterns and breeding territory philopatry. Ph.D. disserta- 
tion, University of Minnesota, St. Paul. 

Gostomski, T. J., and D. C. Evers. 1998. Time-activity budget 
for Common Loon, Gavia immer, nesting on Lake Superior. 
Canadian Field-Naturalist 112: 191-197. 

McIntyre, J. W. 1975. Biology and behavior of the Com- 
mon Loon (Gavia immer) with reference to its adaptabil- 
ity in the man-altered environment. Ph.D. dissertation, 
University of Minnesota, Minneapolis. 

McIntyre, J. W. 1988. The Common Loon: Spirit of north- 
ern lakes. University of Minnesota Press, Minneapolis, 
Minnesota. 288 pages. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Paruk, J. D. 2000. Incubating roles and patterns in Com- 
mon Loons (Gavia immer). Pages 50-54 in Loons: Old 
history and new findings. Edited by J. W. McIntyre and 
D. C. Evers. Proceedings of a Symposium from the 1997 
meeting, American Ornithologists’ Union. North Ameri- 
can Loon Fund, Holderness, New Hampshire. 

Sjolander, S., and G. Agren. 1972. Reproductive behavior 
of the Common Loon. Wilson Bulletin 84: 296-308. 

Taylor, K. 1974. The loon. Adirondack Life 5: 30-37. 


Received 8 November 2004 
Accepted 30 August 2005 


Short-tailed Shrew, Blarina brevicauda, Apparently Killed by Red 


Squirrel, Zamiasciurus hudsonicus 


ROBERT W. NERO 


446 Coventry Road, Winnipeg, Manitoba R3R 1B6 


Nero, Robert W. 2005. Short-tailed Shrew, Blarina brevicauda, apparently killed by Red Squirrel, Tamiasciurus hudsonicus. 


Canadian Field-Naturalist 119(2): 000-000. 


On 2 November 2003, a freshly killed Short-tailed Shrew found beneath a bird feeder was likely the victim of an aggressive 
locally-resident Red Squirrel. The minimal amount of snow cover at the time is hypothesized to have prevented an escape 
such as had been observed in an encounter between these species in a previous year. 


Key Words: Short-tailed Shrew, Blarina brevicauda, Red Squirrel, Tamiasciurus hudsonicus, aggression, Manitoba. 


In mid-afternoon on 2 November 2003, I found a 
freshly-killed Short-tailed Shrew, Blarina brevicauda, 
lying on the seed-strewn ground beneath one of our 
bird feeders which 1s fastened to the trunk of a large 
oak tree. I could hardly have failed to notice the shrew 
had it been there earlier as the site is readily visible 
from out living-room windows. The shrew was lying 
on bare grass which two days earlier I had shovelled 
clear of a thin layer of new snow. Up to this time, I had 
not noticed any shrew sign. The dead shrew, a sur- 
prisingly thin female, had fresh bloody bite marks on 
the back of its neck, on one shoulder, and on the side 
of its head. 

As I could think of no other explanation, I presumed 
that the shrew, which was possibly attracted by the bird 
seed, had been attacked and killed not long before I 
returned home from a short outing, as it was still bleed- 
ing. I guessed that the killer was one of our two resi- 
dent Red Squirrels. One, in particular, an aggressive 
creature that established residence here in the previous 
spring, claimed this oak tree as one of its primary sites, 


regularly driving away other Red Squirrels and Grey 
Squirrels (Sciurus carolinensis); it also frequently 
harassed birds of nearly all species. Nevertheless, part- 
ly because we have two other nearby bird-feeding sites, 
songbirds do regularly visit here. 

Previous observations of a Red Squirrel (not this 
particular one) showing similar belligerence have been 
of some interest. In this same general area, Nero (1996) 
noted that in early December 1995 a Red Squirrel was 
observed attacking a Short-tailed Shrew. That shrew 
struggled fiercely and then escaped its attacker by div- 
ing into a burrow in the snow. In the 2003 incident, 
owing to the minimal amount of snow in the vicinity, 
the shrew had no escape route. 


Literature Cited 

Nero, R. W. 1996. Red Squirrel, Jamiasciurus hudsonicus, 
Short-tailed Shrew, Blarina brevicauda, brief interaction. 
Canadian Field-Naturalist 110: 712. 


Received 18 December 2004 
Accepted 17 April 2005 


2005 NOTES 457 


Extension de I|’aire de distribution de la Couleuvre a collier, Diadophis 
punctatus edwardsii, dans \’est du Québec 


JEAN-FRANCOIS DESROCHES! et BENOIT ROUSSEL? 


! Cégep de Sherbrooke, Département des Techniques d’écologie appliquée, 475 du Parc, Sherbrooke, Québec JIK 4K1, 
Canada 
2 Parc national du Saguenay, 91 Notre-Dame, Riviére-Eternité, Québec GOV 1P0 Canada 


Desroches, Jean-Francois, et Benoit Roussel. 2005. Extension de |’ aire de distribution de la Couleuvre a collier, Diadophis 
punctatus edwardsii, dans lest du Québec. Canadian Field-Naturalist 119(3): 457-458. 


Deux Couleuvres a collier adulte (Diadophis punctatus edwardsi1) ont été trouvées au parc national du Saguenay, dans | est 
du Québec, en 2004 et 2005. Ces mentions se retrouvent a prés 150 kilometres au nord-nord-est de la mention la plus proche 
de l’espece. L’habitat est constitué de forét mixte prés d’escarpements rocheux, dans des vallées. I] pourrait s’agir de popula- 
tions isolées de celles retrouvées plus au sud. 


Two Ringneck Snakes (Diadophis punctatus edwardsii) was found in the Saguenay National Park, in eastern Québec, in 
2004 and 2005. These records are about 150 kilometers north-north-east from the nearest previous record. The habitat is 
mixed forest, with rocky cliffs, in valleys. They may represent isolated populations from those found in southern Quebec. 


Mots-Clés: Couleuvre a collier, Diadophis punctatus edwardsii, aire de distribution, population isolée, habitat, Québec. 


XX 


La Couleuvre a collier (Diadophis punctatus ed- 
wardsil) est une espece discrete qui a historiquement 
été peu rapportée au Québec. Jusqu’en 1990 elle y était 
la couleuvre la moins souvent observée, les mentions 
de l’espece étant confinées au sud de la province (Bider 
et Matte 1991). Il s’agit d’une espeéce discréte et aux 
moeurs fouisseuses, qui estive durant les périodes 
chaudes et séches (Ernst et Ernst 2003). Bien que dif- 
ficile a apercevoir, l’espéce peut étre abondante par 
endroits (Cook 1984; Ernst et Ernst 2003) et pourrait 
étre plus répandue qu’on le croit (Desroches et Rod- 
rigue 2004). Au Québec, la mention de Couleuvre a 
collier la plus a l’est, sur la cote nord du fleuve Saint- 
Laurent, a été faite a Saint-Joachim prés du Cap-Tour- 
mente (Provancher 1874), a quelques 40 kilometres au 
nord-est de la ville de Québec. 

Le 22 juillet 2004, un spécimen adulte a été décou- 
vert dans le parc national du Saguenay, municipalité 
de Riviére-Eternité, canton de Hébert (48°17'35"N; 
70°20'38"). Sa longueur totale est d’environ 27,5 cm. 
II porte sous le ventre une série de points noirs, au cen- 
tre de plusieurs des écailles ventrales. La couleuvre a 
été trouvée morte en bordure d’un sentier, et son état 
laissait présager qu’une faucheuse I’avait découpée en 
morceaux lors de travaux d’entretien. Le spécimen est 
conservé dans la collection herpétologique du Musée 
canadien de la nature (CMNAR-35820). L’habitat est 
une forét mixte dans une vallée encaissée, ol poussent 
le Sapin baumier (Abies balsamea), le Bouleau jaune 
(Betula alleghaniensis), l’Erable a sucre (Acer saccha- 
rum), le Fréne noir (Fraxinus nigra) et des épinettes 
(Picea sp.). La strate arbustive se compose, outre les 
jeunes arbres des espéces cités précédemment, de 
PErable a épis (Acer spicatum), de VIf du Canada 
(Taxus canadensis) et du Noisetier a long bec (Cory- 
lus cornuta). 


FIGURE |. Répartition de la Couleuvre a collier au Québec et 
sur le territoire adjacent. Les mentions faites au parc 
du Saguenay sont répresentées par les points noirs 
numérotés (1 = Riviére-Eternité; 2 = Sacré-Cceur). 
(carte modifiée d’apres Cook 1984; Bider et Matte 
1994; Desroches et Rodrigue 2004). 


Le 27 aoat 2005, un second spécimen de Couleuvre 
a collier adulte a été découvert dans le parc du Sague- 
nay, municipalité de Sacré-Coeur, canton d’ Albert 
(48°15'53"N; 69°56'49"0). D’une longueur totale d’en- 
viron 34,0 cm, il ne porte aucun point noir au centre 
des écailles ventrales. Comme ce fut le case en 2004, 
cette couleuvre était morte lors de sa découverte. Elle 
gisait sur une route d’asphalte, probablement écrasée 
par une automobile. Le spécimen a été récolté et déposé 
au Musée canadien de la nature (CMNAR-35821). 
Lhabitat a cet endroit est une forét mixte, ou |’on 
retrouve l’Epinette noire (Picea mariana), le Sapin 
baumier, des peupliers (Populus sp.), le Bouleau blanc 
(Betula papyrifera), \e Pin rouge (Pinus resinosa) et 
le Pin gris (Pinus banksiana). Beaucoup d’épines de 
coniferes jonchent le sol. 


458 


Ces deux mentions de Couleuvres a collier se retrou- 
vent respectivement a 140 kilometres et 148 kilometres 
au nord-nord-est de la mention connue la plus proche 
de l’expéce (Saint-Joachim (Provancher 1874)) (figure 
1). Elles ont été faites de part et d’autre de la riviére 
Saguenay, soit a l’ouest (Riviére-Eternité) et a lest 
(Sacré-Cceur) et sont séparées entre elles de 30 kilo- 
metres. En effet, la répartition de la Couleuvre a collier, 
dans le sud du Québec, est associée aux différents do- 
maines de |’ érabliere et a celui de la sapiniere a Bouleau 
jaune. La saison de croissance y dure de 160 a 190 jours 
(Robitaille et Saucier 1998). La présente mentions faites 
dans le parc national du Saguenay, se situe dans le 
domaine de la sapiniere a Bouleau jaune, et la saison 
de croissance dure de 140 a 170 jours (Robitaille et 
Saucier 1998). L’Erable a sucre, et dans une moindre 
mesure le Bouleau jaune et le Fréne noir, poussent 
dans le sud du Québec et de maniére isolée au Sague- 
nay (voir les cartes présentées dans Farrar 1996). La 
répartition de la Couleuvre a collier pourrait étre 
semblable. 

Le territoire séparant le Saguenay de la région de 
Québec (mention de Provancher 1874) est tres dif- 
férent. I] est constitué de hautes collines et de monts, 
et fait partie de la zone des sapiniéres 4 Bouleau blanc, 
4 Epinette blanche (Picea glauca), a Epinette noire et 
de la pessiere a mousse. La saison de croissance dure de 
130 a 160 jours (Robitaille et Saucier 1998). Ce terri- 
toire semble peu propice a |’établissement de la Cou- 
leuvre a collier. Nos recherches effectuées dans ce sec- 
teur ne nous ont permis d’y recenser que la Couleuvre 
rayée (Thamnophis sirtalis), laquelle se retrouve au 
Québec presque jusqu’a la latitude 54° nord (MacCul- 
loch et Bider 1975; Bider et Matte 1994; J.-F. Des- 
roches, données non publiées). 

La présence de la Couleuvre a collier au Saguenay 
est trés intéressante; il s’agit d’une extension d’aire 
importante vers le nord-est pour |’ espéce, et possible- 
ment d’une population isolée. Les populations isolées 
de certains reptiles revétent une importance particu- 
lire en conservation (voir dans Seburn et Seburn 
2000). Tout comme Bleakney (1958) l’avait proposé, 
il est possible que le climat plus clément retrouvé dans 
la vallée de la riviere Saguenay favorise certaines es- 
peces de l’herpétofaune qui sont absentes du terri- 
toire environnant, lequel est plus élevé en altitude. 

Il se pourrait également que la répartition de la 
Couleuvre a collier soit plus continue et suive une 
mince bande le long de la céte nord du fleuve Saint- 
Laurent, entre la région de Québec et le Saguenay, 
comme pour les espéces d’ arbres nommeées précédem- 
ment (Farrar 1996) et conformément a la limite de la 


THE CANADIAN FIELD-NATURALIST 


Vol 119 


zone herpétofaunique # 4 proposée par Bleakney (1958, 
page 74). Des recherches supplémentaires sont néces- 
saires afin de préciser la distribution de cette couleuvre 
au Québec, particuliérement dans le nord et l’est, no- 
tamment aux endroits isolés ot les conditions sont 
favorables a l’espéce. 


Remerciements 

Les auteurs remercient Rémi Bouchard pour la dé- 
couverte du spécimen, de méme que Francis R. Cook, 
Daniel Pouliot, Benoit Couture, Hugo Royer (parc 
national du Saguenay) et Isabelle Picard pour leurs 
commentaires sur la version préliminaire du texte. 


Littérature citée 

Bider, J.-R., et S. Matte. 1991. Atlas des amphibiens et rep- 
tiles du Québec 1988-1989-1990, version détaillée. Société 
d’ histoire naturelle de la vallée du Saint-Laurent et minis- 
tere du Loisir, de la Chasse et de la Péche du Québec. 
Québec. 429 pages. 

Bider, J.-R., et S. Matte. 1994. Atlas des amphibiens et 
reptiles du Québec. Société d’histoire naturelle de la vallée 
du Saint-Laurent et ministere de Environnement et de la 
Faune du Québec, Direction de la faune et des habitats. 
Québec. 106 pages. 

Bleakney, J. S. 1958. A zoogeographical study of the amphib- 
ians and reptiles of Eastern Canada. National Museum of 
Canada, Bulletin (155), Biological Series (54). 119 pages. 

Cook, F. R. 1984. Introduction aux amphibiens et reptiles du 
Canada. Musée national des sciences naturelles, Musées 
nationaux du Canada. Ottawa, Canada. 211 pages. 

Desroches, J.-F., et D. Rodrigue. 2004. Amphibiens et rep- 
tiles du Québec et des Maritimes. Editions Michel Quintin, 
Waterloo, Québec, 288 pages. 

Ernst, C. H., et E. M. Ernst. 2003. Snakes of the United 
States and Canada. Smithsonian Institution, Etats-Unis, 
668 pages. 

Farrar, J. L. 1996. Les arbres du Canada. Corporation des 
Editions Fides et Service canadien des foréts, Ressources 
naturelles Canada, Saint-Laurent, Québec, 502 pages. 

MacCulloch, R. D., et J. R. Bider. 1975. New records of 
amphibians and Garter snakes in the James Bay area of 
Quebec. Canadian Field-Naturalist 89: 80-82. 

Provancher, L. 1874. Faune canadienne: les reptiles. Le 
Naturaliste canadien 6: 353-370. 

Robitaille, A., et J. - P. Saucier. 1998. Paysages régionaux 
du Québec méridonal. Les Publications du Québec. 213 
pages + carte. 

Seburn, D., et C. Seburn. 2000. Conservation priorities for 
the amphibians and reptiles of Canada. For World Wildlife 
Fund Canada and Canadian Amphibian and Reptile Con- 
servation Network. 92 pages. 


Received 27 October 2004 
Accepted 24 August 2005 
Revised 4 October 2005 


2005 


NOTES 


459 


Do Juvenile Nearctic River Otters (Lontra canadensis) Contribute to 


Fall Scent Marking? 


ZACH H. OLSON, SADIE S. STEVENS, and THOMAS L. SERFASS 


Frostburg State University, Department of Biology, 101 Braddock Road, Frostburg, Maryland 21532 USA 


Olson, Zach H., Sadie S. Stevens, and Thomas L. Serfass. 2005. Do juvenile Nearctic river otters (Lontra canadensis) con- 
tribute to fall scent marking? The Canadian Field-Naturalist 119(3): 457-459. 


We present photographic evidence in support of the hypothesis that juvenile Nearctic River Otters (Lontra canadensis) con- 


tribute to the observed fall peak in scent marking. 


Key Words: Lontra canadensis, Nearctic River Otter, scent marking, seasonal variation, Pennsylvania. 


Otters scent mark by defecating, urinating, or releas- 
ing anal glandular secretions at conspicuous shoreline 
locations called latrine sites (Melquist and Hornocker 
1983; Swimley et. al 1998). Seasonal variation in 
scent marking intensity has been reported for Nearctic 
River Otters (Lontra canadensis) with peaks in spring 
(March-April) and fall (September-November) in 
Pennsylvania (Serfass 1994; Carpenter 2001; Mills 
2004). There have been many proposed explanations 
for seasonally variable marking intensities in otters. 
Seasonal variation was associated with dominance and 
related to the dispersal of juveniles for a population of 
European Otters (Lutra lutra) in Wales (Macdonald 
and Mason 1987). Carpenter (2001) attributed the 


spring peak in Pennsylvania to mate attraction during 
the breeding season. Similarly, Mills (2004) believed 
increased spring marking was related to advertisement 
of location and breeding condition. 

Home range studies provide evidence in support of 
breeding advertisement and breeding-range defense 
as causes for a spring peak in scent marking (Melquist 
and Hornocker 1983; Reid et al. 1994; Spinola 2003). 
Spinola (2003) reported a spring increase in home 
range and interactions with females by male River 
Otters in a reintroduced population in western New 
York. He invoked two hypotheses to explain River 
Otter spacing patterns: Sandell’s hypothesis (Sandell 
1989) and the resource dispersion hypothesis (Mac- 


FIGURE |. Photograph from the Youghiogheny River (Maryland) latrine site of two juvenile River Otters with one adult. The two 
juveniles are indicated with arrows. The juvenile in the foreground (trailing) appears large in comparison to the other 
River Otters because of its close proximity to the camera. The image is dated 30 June 2004. 


460 


donald 1983; Carr and Macdonald 1986), both of which 
predict female intra-sexual territoriality and large male 
home ranges during the breeding season (Spinola 
2003). Increased scent marking during this period of 
increased movement would likely enhance the effi- 
ciency by which males and females are able to locate 
one another for breeding opportunities. 

Although the possible link between spring breeding 
advertisement and the observed spring peak in scent 
marking has been established, there is a paucity of 
literature examining causes of the fall peak in scent 
marking. Carpenter (2001) and Mills (2004) suggest- 
ed the mechanism for a fall peak in North America 
was an increase in the density of marking individuals 
as juvenile River Otters began traveling with their 
mothers (juveniles were <1 year old). However, we 
are aware of no data that support this hypothesis. 

As part of an ongoing behavioral study of River Ot- 
ters we placed remote, 35-mm cameras (TrailMaster®, 
Goodson and Associates, Inc., Lenexa, Kansas, USA) 
at latrine sites along Tionesta Creek in north central 
Pennsylvania (41°35'N, 78°15'W) and the Youghio- 
gheny River in western Maryland (39°34'N, 79°25'W), 
United States. Two cameras captured what we presume 
to be two family groups (an adult female and two cubs 
as defined by Melquist and Hornocker 1983) of River 
Otters visiting latrine sites beginning 30 June 2004. 
We based this presumption on a size disparity apparent 
among individuals in three photographs containing all 
family group members. The photographs originated 
from two separate latrine sites, one from a Tionesta 
Creek latrine site (15 August 2004) and two from a 
Youghiogheny River latrine site (30 June 2004 and 31 
July 2004). We continued to obtain photographs at those 
latrine sites (n = 2 at Tionesta Creek and n = 10 at the 
Youghiogheny River) of what we assumed to be the 
same family groups through fall and winter 2004. 

To estimate the lengths of the presumed juvenile 
and adult River Otters we took reference photographs 
of both latrine sites. The reference images were taken 
from the same camera mount and position as the origi- 
nal photographs and incorporated meter sticks placed 
at the locations of the River Otters in the original pho- 
tographs. We digitally superimposed each reference 
image onto the original image containing the presumed 
family group of River Otters, thus facilitating an ap- 
proximation of each individual’s total length (tip of 
nose to tip of tail). The 30 June 2004 photograph from 
the Youghiogheny River was not subjected to these 
measurements because the size disparity was readily 
apparent (see Figure 1). In both of the other photo- 
graphs two presumed juveniles were visible, but only 
one was captured completely within the photograph 
frame and could be measured. The presumed adult and 
juvenile at Tionesta Creek were 78.2 cm and 61.4 cm, 
respectively, which yielded an adult length to juvenile 
length ratio of 1:0.79. At the Youghiogheny River, the 
presumed adult was 86.2 cm and the presumed juve- 
nile was 63.8 cm, which yielded a ratio of 1:0.74. The 


THE CANADIAN FIELD-NATURALIST 


Voln ito 


shorter calculated lengths than those expected from 
physical measurements of the same River Otters was 
probably due to the animated posture of the photo- 
graphed individuals. 

Our calculated ratios are consistent with the find- 
ings of Stephenson (1977), who reported November- 
March trapped <1 year-old River Otters in Ontario of 
significantly lower weight and length than all other age 
classes. Merriam (1884) reported observations of young 
otters with their mother in summer and fall in the north- 
eastern United States. Also, Park (1971) listed fall as 
the season juveniles began traveling with their mothers. 
We propose that our photographs were of River Otter 
family groups, juveniles visiting latrine sites with an 
adult, and that the increased number of River Otters 
at latrine sites can explain an increase in the number 
of scats at latrine sites. 

Liers (1951) and Melquist and Hornocker (1983) 
have described River Otter scent marking as a sort of 
positive feedback loop — one otter’s scent marking 
induced other nearby individuals to mark, which in- 
duced another round of marking by the first otter, and 
so on. This type of release stimuli — possibly visual 
and olfactory — has been described for other scent- 
marking carnivores (see Beckoff 1979; Muckenhirn 
and Eisenberg 1972; Peters and Mech 1975; Seiden- 
sticker et al. 1973). Family groups of River Otters vis- 
iting latrine sites would, therefore, leave more scats 
than single otters. Although release stimuli could ex- 
plain increased scat numbers at latrine sites in the fall, 
the function of scent marking as a group has not been 
addressed for River Otters. Ewer (1973) proposed 
that group scent marking in social carnivores could 
facilitate group bonding (page 250). The social sig- 
nificance of scent marking has been described in canids 
(Rothman and Mech 1979), felids (Seidensticker et 
al. 1973), and other mustelids (Buesching et al. 2003). 

Although more information is needed before making 
final conclusions, we believe evidence from the liter- 
ature that a family group begins to travel as a unit dur- 
ing summer, and our observation of juveniles accom- 
panying an adult to latrine sites beginning in late June, 
lends support to the hypothesis that juvenile River 
Otters contribute to increased scent marking in fall. 
Specific aspects of River Otter scent marking at latrine 
sites that have not been addressed in the literature 
include; the composition of marking individuals, the 
periodicity of marking by individuals, and the rela- 
tionship between functional scent marking and elimi- 
nation of feces. 


Acknowledgments 

A United States Fish and Wildlife Service State 
Wildlife Grant (administered by the Pennsylvania 
Game Commission [PGC]), the Pennsylvania Wild 
Resource Conservation Fund (WRCF; nongame tax- 
checkoff), and the United States Department of Agri- 
culture Allegheny National Forest (ANF) provided 
primary funding and support for this project. We are 


2005 


grateful to A. Hayden (deceased) and J. Hassinger of 
the PGC, F. Felbaum and R. Stanley of the WRCF, 
and B. Nelson of the ANF for their cooperation and 
support of River Otter research in Pennsylvania. 


Literature Cited 

Beckoff, M. 1979. Ground scratching by male domestic 
dogs: a composite signal. Journal of Mammalogy 60: 
847-848. 

Buesching, C. D., P. Stopka, and D. W. Macdonald. 2003. 
The social function of allo-marking in the European badger 
(Meles meles). Behaviour 140: 965-980. 

Carpenter, C. P. 2001. Scat marking and use of spraint sites 
by river otters along Tionesta Creek, northwestern Pennsy]- 
vania. M.S. thesis, Frostburg State University, Frostburg. 
95 pages. 

Carr, G. M., and D. W. Macdonald. 1986. The sociality of 
solitary foragers: a model based on resource dispersion. 
Animal Behaviour 34: 1540-1549. 

Ewer, R. F. 1973. The carnivores. Cornell University Press, 
Ithaca, New York, USA. 494 pages. 

Liers, E. E. 1951. Notes on the river otter (Lutra canadensis). 
Journal of Mammalogy 32: 1-9. 

Macdonald, D. W. 1983. The ecology of carnivore social 
behaviour. Nature 301: 379-384. 

Macdonald, S. M., and C. F. Mason. 1987. Seasonal mark- 
ing in an otter population. Acta Theriologica 32: 449-462. 

Melquist, W. E., and M. G. Hornocker. 1983. Ecology of 
river otters in west central Idaho. Wildlife Monographs 
47: 1-60. 

Merriam, C. H. 1884. The mammals of the Adirondack re- 
gion, northeastern New York. L. S. Foster Press, New York, 
New York, USA. 316 pages. 

Mills, M. A. 2004. Scat-marking by river otters in Pennsyl- 
vania and Maryland. M.S. thesis, Frostburg State Univer- 
sity, Frostburg. 178 pages. 


NOTES 


461] 


Muckenhirn, N. A., and J. F. Eisenberg. 1972. Home ranges 
and predation of the Ceylon leopard (Panthera pardus 
fusca). Pages 142-175 in The world’s cats, volume |. Edited 
by R. Eaton. World Wildlife Safari, Winston, Oregon, USA. 

Park, E. 1971. The world of the otter. J. B. Lippincott Co., 
Philadelphia, Pennsylvania, USA. 159 pages. 

Peters, R., and L. D. Mech. 1975. Scent-marking in wolves. 
American Scientist 63: 628-637. 

Reid, D. G., S. M. Herrero, A. C. H. Reid, and T. E. Code. 
1994. Spacing, movements, and habitat selection of the 
river otter in boreal Alberta. Canadian Journal of Zoology 
72: 1314-1324. 

Rothman, R. J. and L. D. Mech. 1979. Scent-marking in 
lone wolves and newly formed pairs. Animal Behavior 
27: 750-760. 

Sandell, M. 1989. The mating tactics and spacing patterns of 
solitary carnivores. Pages 164-182 in Carnivore behavior, 
ecology, and evolution. Edited by J. L. Gittleman. Cornell 
University Press, New York, New York, USA. 620 pages. 

Seidensticker, J. C., M. G. Hornocker, W. V. Wiles, and J. 
P Messick. 1973. Mountain lion social organization in 
the Idaho Primitive Area. Wildlife Monographs 35: 1-60. 

Serfass, T. L. 1994. Conservation genetics and reintroduction 
strategies for river otters. Ph.D. Dissertation, Pennsylva- 
nia State University, University Park. 183 pages. 

Spinola, R. M. 2003. Spatio-temporal ecology of river otters 
translocated to western New York. Ph.D. dissertation, The 
Pennsylvania State University, University Park. 115 pages. 

Stephenson, A. B. 1977. Age determination and morpho- 
logical variation of Ontario otters. Canadian Journal of 
Zoology 55: 1577-1583. 

Swimley, T. J., T. L. Serfass, R. P. Brooks, and W. M. Tzil- 
kowski. 1998. Predicting river otter latrine sites in Penn- 
sylvania. Wildlife Society Bulletin 26: 836-845. 


Received 15 March 2005 
Accepted 7 July 2005 


Book Reviews 


ZOOLOGY 


Damselflies of Alberta: Flying Neon Toothpicks in the Grass 


By John Acorn. 2005. University of Alberta Press, Ring House 
2, Edmonton, Alberta T6G 2E1 Canada. 156 pages. Can 
$29.95. 


I have a very strong interest in entomology, but no 
particular interest in Odonates, save for collecting the 
odd one for my odonatologist colleague. This small 
book has served to enlighten me and further my inter- 
est; I will look at damselflies more closely now. 

In the bulk of the Preface, Acorn waxes about the 
restraints of scientific writing and laments about how 
dull it is to him. He prefers to write “in a frank and 
personal fashion, admitting to one’s influences, desires, 
uncertainties and dislikes.” However, much of “his” 
style (frank, admitting to influences and uncertainties) 
is the basis of scientific writing as well. Literature 
must be directed to an audience, and J don’t think an 
odonatologist would find “dull” the scientific writing 
directed to his or her guild. A quick look at the bibli- 
ography reveals that the vast majority of works con- 
sulted for Acorn’s book is from the scientific literature. 

Having read the book, and having seen episodes of 
Acorn’s TV show and his video interpretations at the 
Tyrell Museum, I believe Acorn is a wonderful author 
for children (e.g. “A big, mean, scary female water 
spider” is part of one caption in this damselfly book), 
but I don’t think that writing for children alone is the 
intended extent of his scope. Acorn’s goal is to write 
for the general public and to get people interested in 
natural history, and this book helps in that. However, 
the author does not give credit to people’s intelli- 
gence...he changes names of structures on the dam- 
selflies simply because they sound too much like jar- 


BOTANY 


Flower Guide for Holiday Weekends 
By E. Larsen and B. Roots. 2005. National Research Council 

Canada, 1200 Montreal Road, Ottawa, Ontario K1A OR6 

Canada. vii + 149 pages. Paper. 

This book is arranged to cover three key holiday 
weekends. The choice of weekends works equally well 
for both United States and Canada [Victoria Day/ 
Memorial Day, Canada Day/Independence Day, Civic 
Holiday/Labour Day]. This is a good concept that will 
be most helpful to novices and visitors. By choosing 
your date you can readily see which of the common, 


gon to him. I’m at a personal crossroads on this point 
—I firmly applaud Acorn’s message of natural history 
education for everyone, but to me, his writing style 
actually detracts from getting the most out of the 
information in the book. For example, I did not find 
the limericks at the beginning of each species descrip- 
tion at all interesting, humorous or otherwise worth the 
space. I believe one must write appropriately for the 
intended audience; there is no such thing as a book 
for all ages. And I do not enjoy reading books that seem 
to be written for children. 

Technically, the book has very good points, as well 
as a few unfortunate oversights. The book is of a con- 
venient size, suitable for taking in the field. The pho- 
tography, while not stunningly appealing, is still quite 
vivid. It is not clear why there is a photograph of a 
dragonfly nymph in the introductory chapter, and none 
of a damselfly nymph; the latter ultimately does appear 
later in the book. Acorn describes the necessity of 
using a 10x lens to be able to identify damselflies in 
the field, yet there are no photographs of any of the 
features at this magnification in the book (there are 
some useful line drawings at the back of the book). 
However, this small book is packed with a lot of use- 
ful information on identification, ecology and behav- 
iour of Alberta’s damselflies, and for that reason alone 
would make a worthwhile addition to a naturalist’s 
library. 


RANDY LAUFF 


Department of Biology, St. Francis Xavier University, Anti- 
gonish, Nova Scotia B2G 2W5 Canada 


showy flowers are in bloom and thus narrow your 
search for the identity of a flower. 

Each species is covered by a good strong narrative 
in layman’s terms. While this text is non-scientific it 
is enlightening and helpful. This is accompanied by 
one to four photos of the plant, its flowers, seed, etc. 
The photos are top quality and are artistically set in 
the book, adding to its charm. The edges of the pages 
are coloured yellow, orange or blue to make it easy to 
pick the weekend. The book is relatively small so 
will slip easily into a pack or pocket. 


462 


2005 


Although 64 flower species are covered, eight of 
these are repetitions. That is, the text and photos are 
virtually identical. I do not see the point of this. A list 
of plants with overlapping flowering times would have 
been sufficient. In the extra space the authors could 
have added some new plants. 


Orchids of Manitoba 


By D. Ames, P. B. Acheson, L. Heshka, B. Joyce, J. Newfeld, 
R. Reeves, E. Reimer, and I. Ward. 2005. Native Orchid 
Conservation Inc., 117 Morier Avenue, Winnipeg, Mani- 
toba Canada. $17.95 + shipping charges. 


This 158-page book with soft covers includes 36 
orchid species known to occur in the province of Man- 
itoba. It contains 218 absolutely beautiful colour pic- 
tures of different aspects of these delightful orchid plants 
in the text plus two more on the front and back covers. 

The text begins with acknowledgements to all the 
individuals who have provided suggestions towards 
the book’s production, the Manitoba Conservation staff 
for their guidance and the use of their maps, Kromar 
Printing for their assistance and getter the colours just 
right, and the financial support provided by the Prov- 
ince of Manitoba Special Conservation Fund, Shell 
Environmental Fund, and The Winnipeg Foundation. 
It is also noted here that the Native Orchid Conserva- 
tion Inc., the publisher of this book, is a non-profit 
organization formed in 1998 to protect unique mini- 
ecosystems and their plant communities. This is fol- 
lowed by “A Brief History of Orchids,” “Conservation 
and Biology,” “Protection of Species and Ecosystems”, 
“Orchid Biology”, “Orchid Habitat” (including a map 
of Ecozones of Manitoba and sixteen pictures of habi- 


ENVIRONMENT 


Book REVIEWS 


463 


This book would make a delightful gift for a traveller, 
cottage owner or those who simply like a good walk. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 
K1J 6K5 Canada 


tats), a “Key to the Orchids of Manitoba” (when in 
flower), and an “Introduction to the Species Accounts.” 
The latter provides information on scientific names, 
abundance, habitat, flowering time, description, aids 
to identification and range maps. 

Pages 45 to 145 are devoted to the illustrations, the 
distribution maps, and the most interesting descriptive 
text for the 36 orchid species treated in this book. Each 
descriptive page provides information on the common 
and scientific names, abundance, habitat, flowering time, 
descriptive information, aids to identification, and addi- 
tional comments. The flowering times for Manitoba 
orchids are presented in alphabetical order of the sci- 
entific names with the common names in brackets on 
pages 146 to 148. This is followed by a bibliography 
on pages 149 to 151 and a Glossary on pages 152 to 
155 and an Index on pages 156 to 157 which contains 
scientific names, common names and selective descrip- 
tive names to help the readers. In addition, a “Fore- 
word” was provided by Paul M. Catling. 


WILLIAM J. CoDY 


National Program on Environmental Health, Agriculture 
and Agri-Food Canada, Research Branch, Wm. Saunders 
Building, Central Experimental Farm, Ottawa, Ontario, 
K1A 0C6 Canada 


Environmentalism Unbound: Exploring New Pathways for Change 


By Robert Gottlieb. 2005. MIT Press, Cambridge, Massachu- 
setts, USA. 2001. 287 pages. US$74.25 Cloth, $20.83 Paper. 


Robert Gottlieb is professor of Urban and Environ- 
mental Policy at Occidental College in California, and 
his study area involves a variety of environmental poli- 
cies and practices, mostly in the Los Angeles area. In 
our modern world, where diversity is publicized as the 
omen of political correctness, but where the survival 
of the richest is often the motivating factor, environ- 
mental awareness is reduced to single questions which 
can be handled by one-issue political campaigns. 
Single issues may facilitate winnable environmental 
battles but the strategy can leave other important areas 
without advocates. Gottlieb places the environment 
into the context of the whole sphere of life and shows 
connected parts interacting to create our quality of life- 
style. Unfortunately, as an urban environmentalist, 
his examples more often show how each of the parts 
has been contaminated by different political and com- 


mercial interests and the big picture purposely left as 
a large unexamined grey area. This book is an attempt 
to reconcile many interests and show inroads which 
have already been made and can reasonably expect to 
be made in a number of industries by response to 
human needs, well-planned activism and some cre- 
ative thinking. 

The focus of the book is urban environmentalism 
exclusively in the United States, but as a study of that 
country, it informs us of the actions of environmental 
groups, the battles they fight and the progress they 
have been making. The ideas presented are ones which 
were studied in Los Angeles but the implications for 
change are important for the rest of North America. Is 
there an alternative in the dry cleaning industry to the 
solvent perchloroethylene? Can fresh fruits and veg- 
etables be locally grown and marketed outside of mul- 
tinational grocery and food distribution companies? 
And can the cleaning industry perform their service 


464 


with environmentally-friendly chemicals, just wages, 
and competition which allows independent operators 
to remain active? 

In answering these questions the book includes a lot 
of data presented in a readable narrative telling how 
the system of business and human resources relates to 
the environment, and how the agents of change have to 
be real agents in people’s lives and business interests. 
In our society respect of the environment also has to 
take into account our financial securities and business 
success or failure. Gottlieb gives us details of different 
cultures, situations, and businesses impacting on envi- 
ronmental justice and pollution prevention situations. 

The book is divided into two parts. The first part is 
a discussion on the bounds and restrictions which peo- 
ple feel when confronted with issues of environmental 
justice. Corporate excuses, small companies’ pressures 
to survive and individual citizens’ concerns for cleaner, 
healthier living are all parts of the discussion. All of 
these human situations impose boundaries on action 
proposed by environmental concerns or pollution pre- 
vention schemes. Gottlieb shows us good news too, 
where some positive steps have been taken in urban 
areas and where work is in progress. 

The second part tells stories of three industries which 
are embracing change under the conditions discussed 
in the first chapters, the dry cleaning industry, the jani- 
torial cleaning suppliers and the community’s supply 
of fresh, economical and local food. Gottlieb takes us 
into a history of the dry cleaning industry, its begin- 
nings, growth, present status and the future of the indus- 
try. There has always been potential for pollution in 
that industry and our desire for hygiene has constantly 
been at odds with the very mechanics of maintaining 
the service which we have adopted. On the horizon 
for us, there are potentials for a cleaner service with 
less pollution, and the alternative of more efficient ways 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


of using the same service but with the same kind of 
pollution. 

The providers of office cleaning service have an 
industry which is rampant with low-salaried workers 
using volatile chemicals with little or no training nor 
protection from fumes, skin contact or emergency res- 
ponse. Smaller operators are forced to compete or go 
under and use the same chemicals also without training. 
Illegal aliens, underage and family workers are all in- 
volved to make a small business operate successfully. 
And when the family and the underpaid workers spend 
their wages, they often have to shop in neighbour- 
hoods which are under-served by the large grocery 
chains because their urban area is too poor to support 
a store which will guarantee a supply of fresh nour- 
ishing fruits and vegetables so readily available to the 
more affluent areas. Local urban bylaws will also keep 
open-air markets from operating in poor neighbour- 
hoods, and local growers and suppliers would not be 
able to market independently due to health by-laws or 
zoning bylaws heavily influenced by the same chain 
superstores which will not locate in the poor neigh- 
bourhoods. 

The book has a wealth of information from careful 
scholarship and even has some good news for the 
future. As most of the environmental literature, it shows 
the amount of work which is going on and the lack of 
or limited successes which environmental policies can 
gain. Much more work remains to be done, but the 
story is being told and Gottlieb tells the story well 
holding our interest in each part. 

Jim O’ NEILL 


St. Mark’s College, 5935 Iona Drive, Vancouver, British 
Columbia V6T 1J7 Canada 


Present address: 28718 Five Mile Road, Livonia, Michigan 
48514 USA 


The Love of Nature and the End of the World: The Unspoken Dimensions of Environmental 


Concern 

By Shierry Weber Nicholsen. MIT Press, Cambridge, Mass- 
achusetts, USA. 2002. 199 pages. U.S.$67.50 Cloth, $18.90 
Paper. 


Nicholsen attempts to bring together the thoughts 
and philosophies of great nature writers, teachers and 
thinkers as well as thoughts and writings of artists and 
psychologists to make her points in this work of nature 
study and doomsday warning. The book wants us to 
consider our place in the development of the world, to 
consider and reconsider the place of people within the 
natural world unfolding, and our presence so strongly 
felt in each part of the world. Human presence is an 
essential part of the world rather than opposition to 
the world, and nature does not need to be suppressed 
in order for us to have a fullness of life. But nature has 
to be meaningful to us first, since our capacity to ap- 
preciate the world around us impacts upon our own 


interior life and perception of what will sustain our 
lives. If we cannot see the world around us, we can- 
not care for it. 

The text reads as much like a poem as a discussion 
in many places, with quotes from a great variety of 
writers illuminating the points which Nicholsen tries 
to make. I found myself leafing through the book to 
reread quotes from Thoreau, Paul Shepherd, Gary Sny- 
der, Aldo Leopold and Jack Turner. Many other writers 
from a variety of traditions are included illustrating the 
views of nature synonymous with art, spirituality, phi- 
losophy and psychology. In fact, the Name Index lists 
135 different authors, some of them quoted several 
times, a large number of source authors for a 200-page 
text. The number of authors included shows us the mag- 
nitude of literature supporting Nicholsen’s thesis; love 
of nature is one of the defining aspects of art, spiritu- 
ality and philosophical thinking over the ages. 


2005 


We are faced with a crisis but we have responded 
with apathy, and unconsciously decided not to react to 
the crisis at all. At the same time, everyone we meet is 
has some concern and appreciation for at least part of 
the environment. How does this split in our thinking 
persist? Our society has grown so used to taking the 
world for granted that we can neither be shocked nor 
shamed into doing anything about the ecological cri- 
sis because we are inundated with other more shocking 
news every day and we have made the decision that 
we will continue to exist without paying attention to 
the evidence before our eyes. This decision, expressed 
by psychologist Harold Searles, gave Nicholsen the 
idea for the book. Our decision, Nicholsen wants us 
to know, is wrong and a denial of the evidence we 
can see as well as denying North American Native 
wisdom, Christianity, Islam, and Buddhism among 
other teachings of the ages. Artists, philosophers, and 
teachers of nature all give evidence of what we are 
missing, and what we are destroying, but we do not 


BOOK REVIEWS 


465 


respond because we have already made up our minds 
not to take the wisdom into account. 

The book recognizes that our relationship with our 
world is destructive and has always been so, from the 
moment when we began to kill wildlife and uproot 
plants to nourish ourselves. However, killing and up- 
rooting are only the beginning and not the end of our 
relationship with our world as we know, but could well 
become the fulfilment and fate of the world if we can- 
not see beyond incidentals. This book is an opportu- 
nity to see beyond where we are, which direction we 
seem to be going and an invitation to visit the wis- 
dom of the ages. The sages teach us that the progress 
of nature does not despair of the possibilities of the 
future and neither should we. Nicholsen shows us a 
bleak path of destruction, but with the thread of hope 
that nature itself interpreted by writers past and present 
can lead us back to a positive relationship with the 
world in which we live. hay OM NEnE 


28718 Five Mile Road, Livonia, Michigan 48154 USA 


Survival by Hunting: Prehistoric Human Predators and Animal Prey 


By G. Frison. 2004. University of California Press, Berkeley. 
xix + 266 pages. Can $47.25 Cloth. 


The archaeological excavations in which I have been 
involved in interior western Canada often yield large 
amounts of animal bone, eloquent testimony to the 
importance of hunting for people in the past. Like most 
modern urban people, however, I have never hunted 
or butchered animals and so my ability to understand 
what I see in the archaeological record lacks this source 
of enrichment. In his fine book, Survival by Hunting, 
George Frison argues that direct hunting experience 
is a fundamental source of knowledge for archaeolo- 
gists and laments the fact that most know nothing of 
this activity. He expresses his “dissatisfaction with 
ethnographic and archaeological interpretations of 
human hunters and hunting that fail to acknowledge 
the years of experience and the accumulation of knowl- 
edge of animal behaviour required to become a success- 
ful hunter.” In Chapter 9 (“Concluding Thoughts”’), he 
reiterates his frustration “that human hunting has 
wrongly been viewed as a kind of instinctive behav- 
iour not worthy of serious anthropological study.” On 
the contrary, he states that “‘killing an animal’ hardly 
describes the body of learned behaviour acquired over 
a long period of time that leads to that final act.” He 
argues persuasively for an experiential approach to 
archaeology, using his own life and career as exem- 
plars. As such, this book contains a great deal of infor- 
mation about animal behaviour and biology, though 
viewed from the perspective of a hunter. 

Although he does not express it in quite these terms, 
my reading of Frison’s argument suggests that he is 
impatient with the attitude of students who regard hunt- 
ing and killing an animal as easy, whereas they regard 
the butchering and use of the remains as the difficult 
part requiring interpretation. Perhaps this is because the 


process of hunting, as opposed to the kill and carcass 
use, leaves scant archaeological remains. Certainly, there 
are sites, such as Head-Smashed-In Buffalo Jump in 
southwest Alberta, where drive lanes and terrain con- 
figuration allow some reconstruction of the hunting 
strategy. But in most cases, especially where sites are 
deeply buried, this level of information cannot be recov- 
ered. The hunting process, as Frison describes it, in- 
volves an intimate knowledge of animal behaviour with 
a sophisticated understanding of terrain, and demands 
adaptability, persistence, and considerable strength and 
physical skill. Much of this, therefore, resides in the 
mind of the hunter and not in the material culture or 
discarded faunal remnants found at an archaeological 
site. 

The first two chapters (“Where the Buffalo Once 
Roamed” and “The Education of a Hunter’) draw large- 
ly on Frison’s own life experience. He describes how 
he grew up in northern Wyoming in the 1920s and 
1930s, learning to ride and hunt with his grandfather, 
partly to undertake predator control, and partly to sup- 
plement the food supply on the ranch. He spent much 
of his early adult life as an outfitter and guide. He re- 
counts how his interest in archaeology grew from his 
encounters with sites and artifacts while he was out on 
the land. His fascination with this material and the peo- 
ple of the past who made it increased, until finally, in the 
1960s, he was persuaded to enter the academic world, 
gaining formal training in the field that absorbed him. 
In the decades since then, he has excavated and pub- 
lished accounts of many of the most widely-known 
archaeological sites in the interior northwestern Unit- 
ed States. These include the Casper site, Agate basin 
site, Horner site, and Mill Iron site. Perhaps because he 
was an outdoorsman first and an archaeologist second, 
his work has always included, perhaps to a greater ex- 


466 THE CANADIAN FIELD-NATURALIST 


tent than many other contemporary archaeologists, an 
appreciation for the subtleties of the terrain around 
the sites. 

The heart of the book concentrates on the hunting of 
the large mammals that were major food sources for 
people through the past eleven millennia. The region- 
al focus is the interior of North America, mainly the 
northern Great Plains and adjacent Rockies of the Unit- 
ed States, the areas where Frison has spent most of 
his life and career. Frison begins his survey by con- 
sidering Paleoindian, notably Clovis, hunters and the 
approaches they might have taken to hunting the now 
extinct megafauna, especially mammoths, around 
11 500 to 10 000 years ago. This chapter is more spec- 
ulative than the rest and, because there are no mam- 
moths around to hunt today, it is obviously less based in 
personal experience. Elephants, perhaps, are the near- 
est one can now get to mammoths, though it is unclear 
how closely their behaviour and reactions to hunting 
are analogous. Frison describes some butchering experi- 
ments undertaken during elephant culls in Zimbabwe. 
From his account, cutting up an elephant with a stone 
knife is a lengthy, messy, and tiring process! 

The next three chapters each focus on one of the 
“big three” prey mammals of the northern plains and 
Rockies: Bison, Pronghorn, and Mountain or Big Horn 
Sheep. As merits its importance in the archaeological 
record, the bulk of this discussion (almost 60 pages) 
focusses on Bison. This chapter begins with a consid- 
eration of the palaeontological evidence for different 
Bison species — a subject about which there is still much 
debate — followed by a survey of selected archaeolog- 
ical Bison kill sites. Frison reviews the several strate- 
gies that were used to hunt and kill Bison, resulting 
in kill sites of different types including arroyo traps, 
Bison jumps, and corrals. He then presents some inter- 
esting observations of Bison behaviour based on Bison 
ranching, knowledge that is needed in order to judge 
how the animals might react to various situations. One 
point that comes through strongly from these accounts 
is that cattle are not a good analogue for Bison because 
their behaviour and biology are quite different. Frison 
structures sections on Pronghorn and Big Horn Sheep 
in the same way, with an introduction to the animals’ 
biology, ecology, and behaviour, emphasizing traits, 
such as herding, that would affect hunting strategies. 
He summarizes ethnographic accounts of hunting and 
then describes the archaeological signatures of pro- 
curement. These are often different and characteristic 
for each animal: corrals for Pronghorn and small traps 
or pens made of logs for Big Horn Sheep in the high 
country. 

Chapter 7 deals with the hunting of various other 
animals in the same region. These include large ani- 
mals — specifically deer, both White-tailed and Mule 
deer, Elk, and Black and Grizzly bear — and small 
mammals, especially rodents, and birds. Although 
sometimes present, these tend not to feature as promi- 


Vol. 119 


nently in the faunal assemblages from archaeological 
sites in the regions under consideration. Indeed, Fri- 
son questions whether “small mammal procurement 
should be considered in the category of ‘hunting’”. 
Nevertheless, birds and small mammals may have been 
important diet items in some circumstances, although 
perhaps not as “archaeologically visible” as large mam- 
mals. The discussion of birds is quite brief and deals 
with only two (Sage Grouse and Blue Grouse); I found 
it interesting that there was no discussion of waterfowl 
hunting. Frison comments (page 178) that he has no 
experience with hunting Caribou, another large mam- 
mal that was an important food source for people fur- 
ther north. Frison briefly describes an unusual arrange- 
ment of canid faunal remains. Although this seems 
one implication, he does not discuss potential cere- 
monial aspects to this arrangement, while he does 
mention possible ceremonial aspects to arrangements 
of Elk antlers and Big Horn Sheep horns. However, 
throughout this book, Frison’s focus is on hunting 
animals for food; other possible reasons for animal 
procurement are only touched on tangentially. 

Throughout these chapters, Frison has mentioned 
the various weapons that were used in hunting. Chap- 
ter 8 brings this together and concentrates more explic- 
itly on “Weaponry and Tools Used by the Hunter”. The 
main weapons include the spear, the bow and arrow, 
and the atlatl. Stone tools include those used to butch- 
er and process an animal, such as knives, hammers, and 
scrapers. With respect to these weapons and tools, 
our perceptions may be strongly coloured by what is 
recovered at archaeological sites, that is, mainly arti- 
facts made of stone, although Frison does include some 
discussion of weapons and tools made from other mate- 
rials, such as bone, antler and ivory. There is a further 
bias in the archaeological record because only part of 
the weapon system, the projectile point, is usually recov- 
ered. The other essential parts of the technology, such 
as wooden spear shafts, sinew lashings, bow strings, 
etc., which are made from more perishable materials, 
are rarely preserved in this region, except in unusual 
settings such as dry caves. 

This is a truly fascinating book. It is a “must read” 
for professionals involved in archaeology, and plains 
archaeology especially. But anyone with an interest 
in the lifeways of people in North America prior to 
European settlement will find much to ponder on here. 
For naturalists, the main interest of the book lies in the 
practical observations on behaviour of some major large 
mammals of the continental interior. A career statement 
from one of the towering figures in late 20" century 
North American archaeology, this volume represents 
the distillation of a lifetime’s experience garnered by 
fieldwork and reflection. 


ALWYNNE B. BEAUDOIN 


Royal Alberta Museum, 12845 102 Avenue, Edmonton, 
Alberta TSN OM6 Canada 


2005 


BOOK REVIEWS 


467 


Wildlife, Conservation, and Human Welfare: A United States and Canadian Perspective 


By R. Taber and N. Payne. 2003. Krieger Publishing Com- 
pany, Malabar, Florida. US$29.50 Cloth. 


This book is a firework; it is full of incredible details 
on managed and mismanaged wildlife and natural 
resources. The beauty is that this convincing text comes 
from two acknowledged experts who are actually 
within the wildlife management circles, and who have 
worked in this discipline for over 30 years. Therefore, 
one can hardly find a more honest and matching 
description of the current state of wildlife management 
in North America, and elsewhere. It will hopefully put 
publications by B. Lomborg (The Skeptical Environ- 
mentalist) and others to rest, and lighten up the en- 
trenched old-fashioned sections of the wildlife com- 
munity towards new horizons. The authors state that 
“...the traditional resource coteries tends to resist 
change. It has a high level of internal coherence and 
devotion to a well-defined philosophy, and is led by 
individuals educated in an earlier age”. 

This fascinating book offers 14 chapters showing 
shockingly how far away we still are from a truly sus- 
tainable wildlife management; e.g., as requested by the 
Rio Convention 1992, by common knowledge and by 
textbooks of science-based resource management. The 
Chapters 5 “European Trade” and 6 “Exploration and 
Settlement” expose our recent globalization as a sim- 
ple repeat of history. I suggest Chapter 12 “Broaden- 
ing Conservation and Wildlife” as required reading 
for any scholar of wildlife and natural resource admin- 
istration. 

The authors expose nicely that the widely heralded 
concept of “multiple use” often simply meant nothing 
else than: “Fiber first, and wildlife last”. They pro- 
vide extensive examples of how harmful unsustain- 
able management of resources is: in part, Alaska was 
sold to United States from the Russians due to their 
overexploitation of fur and other resources (oil and 
gas wasn’t known by then). The Roman Empire fell 
due to the failure of blending economy and society, 
including deforestation, overgrazing, erosion and pol- 
lution of the natural environment. Once the Roman 
dominance ended approximately 500 AC, it led into 
the Dark Ages lasting from 400-1400 AC; a so-called 
period of intellectual stagnation. Another example 
referred to in this book: in 1841 the Russians aban- 
doned their southernmost port, Fort Ross, on the coast 
of northern California because the local supply of 
marine mammals (not only sea otters but also fur seals) 
was already exhausted by overhunting. It was only 
the gold findings in California in 1848 that brought a 
new group of settlers into the region. 

During the 218 pages of fascinating text the authors 
destroy the Myth of Superabundance of wildlife in 
North America. The United States list of “vertebrates 
in jeopardy” consists of over 78 birds, 14 reptiles, 12 
amphibians and 71 fishes, and of additional 148 inver- 
tebrates, and 596 plant species. 


The book makes a strong case that the natural envi- 
ronment is a vital component of the human environ- 
ment. It shows nicely that the developed nations have 
22% of the world’s human population, but use 88% 
of worlds’ resources, 73% of its energy, and generate 
most of its waste and pollution. Whereas 78% of the 
world population consumes only 12% of earth re- 
sources. Instead of calling the Third World “less dev- 
eloped”, the authors suggest to call them “less con- 
sumptive” (and consequently the western world “more 
consumptive”). The authors make clear that North 
Americans use the land and energy resources from the 
rest of the world: the average U.S. citizen consumes 
50 times more than average citizens in India. Despite 
subsistence lifestyles in rural areas, it was the people 
of the urban activism, concerned with non-material 
rather than material satisfaction, that started “Earth 
Day” (actually led by U.S. Senator G. Nelson). It result- 
ed eventually into the green or environmental move- 
ment and had a global impact. 

Perhaps some readers would not expect such revo- 
lutionary and “challenge the hierarchy” thinking and 
statements from authors that are Vietnam veterans. But 
these successful Wildlife Managers with an emeritus 
status, one of them a former Aldo Leopold student, 
have gone environmental for a good reason: our current 
economy regime harms wildlife and habitats alike. The 
authors do a brilliant job in summarizing their own 
research work as well as the current wildlife habitat 
dilemma of the second millennium. For instance, one 
reads throughout the book that market incentives fail 
too often to conserve or use biodiversity on a sustain- 
able level, ““...they even facilitate degradation of eco- 
systems and depletion of species”. One of the provided 
examples deals with the Hudson’s Bay Company 
which, clearly held a monopoly, and did what monop- 
olies do best: unconstrained exploitation until they 
overexploited the resource; buying low and selling high 
but without any considerations of long-term sustain- 
ability. History taught their business attitude a lesson. 

The authors provide a brilliant analysis and descrip- 
tion for furbearers, and how it affected the global com- 
munity: already by 1840 Beavers became commer- 
cially extinct, almost world-wide! By then, silk replaced 
fur, and prices dropped for Beaver. In contrast to other 
fur animals such as fox, lynx, sable and ermine, Beavers 
are herbivores and thus they occur in high populations 
and are readily harvestable. The fur trade began approx- 
imately in 1500 in Cape Breton, Canada, delivering 
products to France and Spain/Portugal. Quickly, North 
American natives then became part of a global mar- 
ket economy; e.g., via Holland, England and France. 
Fur trade meant predominantly “beaver”. Before the 
North American fur quest started, Europe, as well as 
parts of Russia, were already hunted out for Beaver 
pelts. The introduction of steel traps in 1750 and their 
mass production did the rest to harm populations of 


468 


Beavers and other furbearers. As authors show, Beavers 
in the New York region, the location of one of the first 
bases of the Hudson’s Bay Company, were quickly 
eliminated. The nearby St. Lawrence and Great Lakes 
regions, down south to the Gulf of Mexico even, were 
next, and then followed by the boreal forest zones. Both 
sides of the Pacific Coast came last, which got en- 
forced by the North West Company and Russian trade 
companies. Once the Beaver was “done”, Muskrat and 
Raccoon were next in line. 

This book is one of the best reads as a resource for 
wildlife management issues and related details: In 
North America, since the 16" century, 94 birds and 
36 mammals became extinct, and 253 birds and 316 
mammals are almost extinct. As the Hawaiian example 
shows, “Captain Cook opened the way to the extinc- 
tion of 90% of the indigenous species of birds...and 
introduced species (870 plants, 2000 invertebrates, 
80 vertebrates)”. Within only 20 years of its discovery 
by the western world, Steller’s Sea Cow became extinct; 
it took approximately 100 years for the Sea Otter. Asian 
Lions were found for thousands of years all over Asia, 
Africa and southern Europe; but in recent times they 
were reduced decade after decade and are now only 
found in India. As further shown, the experiences with 
Galapagos tortoise, Gray and Bowhead Whales are 
not any different. “Among the American colonies, 
local extirpation was the order of the day”’. It’s simply 
part of the ‘American way of life’; one of the largest 
export products ever existed. Settling the United States 
automatically meant extirpation for species such as 
Moose, Elk, Bison, White-tailed Deer, Wild Turkey and 
Beaver. “Wildlife populations in the more densely set- 
tled east were declining by 1850...”. And “... as rail- 
roads connected markets a lively wildlife trade contin- 
ued”. Frontier farmers turned into commercial producers 
with railroad connections increasing pressures on the 
wildlife resource which resulted in further declines 
and extinction. Nowadays, the eastern American Black 
Duck population has declined due to acid rain. 

This books brings interesting native issues to the 
forefront as well: refuges along native tribe borders 
always had enough deer and wildlife because they 
represented an “unused grey-zone”’. But their wildlife 
abundances crashed immediately once tribe borders 
changed. The quest by European’s for furbearers 
brought human diseases into the land previously domi- 
nated by natives, an area over 5000 miles in diameter, 
and one of the worst cases of disease spread ever known 
in human history. Subsequently, that reduced native 
hunting pressure on wildlife; e.g., Bison populations 
sextupled! Nowadays, natives in Canada and elsewhere 
face two options: merge with dominant industrial cul- 
ture, or adapt their traditional culture to new conditions. 

The authors show that federal public lands are under 
direct control of the president of the United States via 
U.S. federal agencies [e.g. U.S. Fish and Wildlife Ser- 
vice (USFWS) and U.S. Bureau of Land Management 
(USBLM)]. However, only 3% of the United States is 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


actually protected (whereas Costa Rica has over 12%). 
Further, “most private land in U.S., more over the 
half total, is managed on economic principles within 
short time frames and with no discernable concern for 
human welfare in other than monetary terms, or for 
ecological sustainability.” 

Very enlightening are also the presented views and 
constraints about the wildlife and management pro- 
fession itself: “A manager in industry who does not 
please the stockholders can be replaced. A manager 
in a public agency who does not please the tradition- 
al agency clientele can be punished in several ways, 
among them transfer, reduction in budget, no promo- 
tion, or salary increase.” But eventually, this person 
is still irreplaceable, blocking progress and contribut- 
ing to the Cult of Incompetence which is nowadays 
widely seen in governmental agencies (“dead wood”). 
As shown in this book, such a situation does not only 
create frustration but supports environmental devas- 
tation on large scales. 

Need an example? The authors provide plenty: “Biol- 
ogists closest to the natural behavior of endangered 
species have encountered the natural behavior of gov- 
ernment agencies and its negative consequences for 
species recovery. In a broader view, it seems that prac- 
tically all human entities involved in an endangered 
species recovery program will benefit most, materially, 
as long as the species does not become extinct but never 
recovers to a viable population level”. The direct effect 
from this entrenched but incredibly harmful manage- 
ment and governmental culture of wasteful nothingness 
is that 65 forms of mammals are in jeopardy (accord- 
ing to the USFWS list from 2002; 251 forms are in 
jeopardy in other countries); many more life forms are 
under consideration but this is way beyond capability 
of the agencies. Despite the governmental management 
and mandate, much wildlife is not adequately consid- 
ered nor managed. 

The authors emphasize “...the ancient continuing 
tendency towards Tribalism” and “testosterone” in the 
wildlife discipline: especially young men follow agen- 
cies and clubs, including their agendas such as pro- 
vided by professors, coop units, USFWS and CWS. 
The authors quote: “... government programs at every 
level are the responsibility of a multitude of separate 
regulatory agencies, each with a primary interest in 
self-preservation and continued customary service to 
its traditional clientele, and steadily supplied with new 
recruits from specialized professional curricula at uni- 
versities.” And “...the cadres managing the various re- 
newable natural resources had inevitably been becoming 
more and more inner-directed, 1.e., out of touch with 
these new cultural changes. Each managerial group was 
recruited from students attracted to the appropriate pro- 
fessional curriculum, the student was indoctrinated with 
the traditional philosophy of that particular resource 
by instructors who had a similar education and had 
often served in the industry or agency dealing with that 
resource. Each renewable resource then had its adher- 


2005 


ents: students, instructors, researchers, agencies, indus- 
tries, and particular user-groups, supporting and served 
by sympathetic elected legislators. Such a cadre focused 
on a particular resource and became more and more 
internally coherent over time, producing accepted terms 
and philosophies of management, with members meet- 
ing one another periodically to re-enforce the mutual 
vision of how their resource should best be managed. 
Eventually, as the whole society developed new per- 
spectives, these traditional professional resource groups 
began to lose public trust and esteem. Perceived as a 
threat to draw each resource cadre together in a defen- 
sive posture.” 

Despite all these great and important contributions, 
I find the book does not address really well the actual 
mechanisms of how wildlife links with humans; e.g., 
why only 7% of North American adults hunt. For 
over 100 000 years humans made 99% of their living 
through hunting, and at least 4 of their diet consisted 
of meat. Taber and Payne show cases where hunting 
contributed to extinction of large mammals, such as 
with the North American native tribes of the Cahokians 
and Hohokam; humans as r-strategists. For the Amer- 
ican sportsman, the loss of prairie grouse (Sharptail, 
Greater Prairie Chicken, Lesser Prairie Chicken) got 
simply compensated by the introduction of alien species 
such as Ring-necked Pheasant and Gray Partridge. The 
authors state that many national leaders were former 
soldiers, and this can affect wildlife due to the hered- 
itary soldier-rule and aristocrat views which got direct- 
ly imposed onto the environment then, and onto its 
legal administration. This forms a huge and lasting 
culture, as can be seen to this very day in the German 
Prussian, French, Russian, Royal English and even 
African hunting codes shaping the current set-up of 
wildlife, habitats and resulting attitudes of the globe. 
The authors make a strong case that the Legal Hunt- 
ing rights for the common citizens in England were 
gradually reduced to nothing. 

The text sections for National and Protected Parks 
are a great and very informative read: “When wildlife 
in the colonies became threatened, generally by human 
population increase and pressure on wildlife habitat 
(as it had in western Europe in medieval times), colo- 
nial administrators set aside some of the best remaining 
habitats as wildlife sanctuaries (just as their ancestors 
had done in medieval times). Most of these were estab- 
lished in 1930, til break up of colonial powers in the 
1950s. Cultural concepts of wildlife conservation came 
to Canada and the U.S. principally from England...”. 
This approach was often encouraged by the upper 
class society; e.g., reflected in many wildlife funding 
schemes. In America, the first National Park came 
into existence in 1872 in Yellowstone (where hunting 
was still allowed for the subsequent 22 years), and in 
Canada in 1887 with the Rocky Mountains park (now 
Banff), whereas the first real reserve was founded as 
early as 1832 with the Arkansas hot springs. In 1881, 
the first U.S. forest reserves got established, followed 


Book REVIEWS 


469 


1906 by the Forest Reserves Act. This is the lasting 
effect when hiring leaders with vision (in this case 
G. Pinchot 1901). 

However, the authors report that despite National 
Parks, many endangered species are found in habitats 
fully shared by humans and far from protected natural 
areas. “No reserve, no matter how large, is large enough 
to sustain a viable population of its more space-demand- 
ing species”. The buffer zones are supposed to improve 
this situation but this is an area where humans en- 
croach, too. Authors hint to the controversial point of 
view of a consumptive use of National Parks in the 
Third World. 

I really like the great descriptions and summaries 
of legal events that put wildlife management in a pol- 
icy context: By 1969 the U.S. just had experienced 
three decades of unparalleled prosperity when the U.S. 
National Environmental Policy Act (NEPA) was ini- 
tiated with the U.S. National Council on Environmen- 
tal Quality (NCEQ) overseeing this process. The intent 
of NEPA is to force agencies predicting effects as far 
as possible in a quantitative fashion to avoid inciden- 
tal damage to environmental values that their mana- 
gerial decisions might cause. NEPA requires major 
things to be considered in regards to the environment. 
“These policy statements invoke the need to fulfill, 
assure, attain, preserve, achieve, and enhance social 
and environmental values in conservation and renew- 
able resources...”. In theory, this provides for a great 
template balancing the economy with social and 
environmental issues. However, “The intent of NEPA 
also was blunted by agency reluctance and inability to 
adapt to new directives and to competition between 
agencies for NEPA-generated funds and tasks”. Sec- 
ondly, land management agencies often had no clue 
about the actual land content as they only managed 
for timber, grass, flood control, military ranges, etc. 
Quantifying biodiversity must always be a priority 
for biodiversity conservation. Lastly, if a controversy 
occurs, each side tries to demand the burden of proof 
from the other side. Obvious legal and administrative 
problems exist with the actual “jeopardy opinion”, 
which results in the acceptance of the actual burden 
of proof. 

Here another statement based on the huge experi- 
ence by the authors with a high academic status that 
does not help to increase public trust in governmental 
actions when it comes to the environment and human 
welfare: “In a culture of public employees, every play- 
er must be aware, for welfare of self and dependants, 
that he/she is vulnerable. The higher people advance 
in the agency, the more they have to lose. On their 
own behalf, then, as well as their belief in the virtue 
of their organization, they will tend to place the wel- 
fare of their organization above any different good”. 

The book certainly presents in detail another global 
milestone in legislation introduced in 1971: The U.S. 
Endangered Species Act (ESA). But progress on recov- 
ery of endangered species is slow due to too many 


470 


bureaucratic hurdles, and due to a slow listing process 
and inadequate funding of the act. The authors make 
it very clear that for listing species in ESA, any eco- 
nomical considerations have to be ignored: it should 
be purely driven by decline and extinction concerns. 
But instead, and often due to financial constraints, right 
now 600 Category I species await listing, 3000 Cate- 
gory II species still await research and sound assess- 
ment “But environmentally ignorant politicians often, 
usually, reduce environmental budgets”, and “The res- 
ponsible services respond by emphasizing work on 
charismatic; 1.e., glamour species ...” to obtain easier 
funds from the politically powerful urbanite. “Fur- 
thermore, by imposing a more rigorous standard of 
review, decisions of often technical scientific issues 
are shifted from an agency with substantial biological 
expertise, to judges who have none”. 

Habitat issues get well-covered, and authors promote 
co-management of the land. They show that it is a huge 
short-coming for wildlife that the United States has an 
ESA but not a Habitat or Ecosystem Act. As history 
showed already in United Kingdom, forest cover loss 
and human pressure resulted in severe declines of 
Aurochs, Forest Bison, Brown Bear, Wolf and Red Deer. 
In North America, Atlantic forests had been kept open 
by natives through the use of fire in November (as 
supported by the well-known fact that the first Euro- 
pean seafarers detected land long before seeing it by 
smell). This type of land management favored the 
Heath Hen, Elk and Bison; but it all changed in an 
evolutionary eye blink with the advent of farmers. In 
the U.S., and due to the extent of the land, trespassing 
was hard to control and a charge for use was impossi- 
ble. Thus, everybody could use the available game. 

From 1982 onwards, the ESA requested that for 
each endangered species a Habitat Conservation Plan 
(HCP) had to be added in order to address critical habi- 
tat. This shifted now the burden of proof to the agency, 
which consequently resulted in only a few completed 
HCPs, so far. 

This publication gives a nice overview of North 
American wildlife management history. Before ESA, 
most attention was given to game species only, since 
agencies were in charge dealing with game manage- 
ment. It is only since 1910 that every U.S. state actu- 
ally has had a wildlife agency. The federal agency did 
not come into place before 1940 (USFWS: 1947 for 
Canadian Wildlife Service CWS). Five periods of 
Wildlife Management are presented: 1600-1849 era 
of abundance, 1850-1899 period of overexploitation, 
1900-1929 period of protection, 1930-1965 game man- 
agement and 1966 to present environmental manage- 
ment. 

The International Wildlife Conservation Chapter I 
find an outstanding read also. The global goal still 
appears to turn everything into sustainable use; failures 
of TRAFFIC and CITES conventions in administer- 
ing this movement are shown. Three goals of interna- 
tional conservation are: (i) preservation of species, 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


(ii) integration of economic development and nature 
conservation, and (111) effective cooperation of rural 
local people in wildlife conservation. Following this 
principle, authors present USAID as a development 
agency that has been heavily involved in hydro power 
projects world-wide. The roles of IUCN, Red Data 
Book, Survival Service Commission, WCMC, WWE 
and IWC get discussed in detail, too. A strong plea is 
made that U.S. should not withdraw from UNESCO 
(an agency that started with help from the English 
zoologist Julian Huxley, and which therefore included 
nature conservation within its scope). 

This book shows that whatever happens in U.S. will 
eventually happen in Canada as well. It also shows 
that Canada is way behind when it comes to Wildlife 
Management, and that it is certainly not world-lead- 
ing; e.g., the Canadian Environmental Protection Act 
got implemented as late as 1993. However, with Cana- 
da following U.S. almost blindly, at least consistency 
is assured across the North American continent; e.g., 
when compared to the diverse, if not even chaotic, 
policies in the European Union. The authors are not 
based in Canada, and some issues presented in the book 
for Canada sometimes appear a little too simplistic. 
Throughout the book, Canada gets portrayed as being 
similar to U.S., whereas U.S. has a much stronger NGO 
pressure than Canada has ever seen and experienced, 
making huge differences between the two countries. 
Despite the claim by the authors, Canada 1s definitely 
not further advanced in GIS and Satellite Imagery than 
U.S. Gust have a look at the GAP programs for in- 
stance). From what I know, the Canadian Ecoregion 
approach to landscapes and management mentioned 
is actually very weak, to say the least. 

This is an environmental history book par excellence 
but, unfortunately, without any relevant graphs, figures 
and maps. I am not a big fan of the reference organi- 
zation either: references are not linked to the text and 
statements, and are hard to connect back from the text. 
Many chapters also have the same references cited sev- 
eral times. In some occasions, the text is a re-expla- 
nation of already published papers. The human wel- 
fare part could be stronger elaborated on, Adaptive 
Management principles by Carl Walters are hardly 
mentioned, nor any modern digital data issues for wild- 
life and habitats. For people with a European Union 
background, some of the related text sections might 
appear imprecise and blurred. At a few text sections, 
I am really at odds with the authors; for instance, they 
suggest that children had to be protected from wild 
predators, and that farming would have had negative 
effects on human life length and quality. Smaller errors 
can be forgiven (Domestication of African Elephants, 
Sperm Whale as the only large whale in tropical waters; 
U.K. being fully representative for Europe). 

As a wildlife practitioner myself, I am extremely 
grateful that these two very experienced authors with 
highest academic ranking devote the book to “... stu- 
dents and field biologists acting under often trying 


2005 


circumstances to strengthen the factual base for sus- 
tained positive relations between human and other 
forms of life.” We need more of this, indeed. One 
might hope from this great book that Wildlife Man- 
agers will read, learn, and become environmentally 
considerate, eventually. But unfortunately, so far, the 
current facts and global political climate are just show- 


NEw TITLES 
+ Available for review * Assigned 


Zoology 


Amphibian Declines: The Conservation Status of United 
States Species. Edited by Michael Lannoo. 2005. University 
of California Press, 2120 Berkeley Way, Berkeley, CA 94704- 
1012. Distributed by NHBS2-3 Wills Road, Totnes, Devon 
TQ9 5XN, UK £62.00 Cloth 


The Amphibians and Reptiles in Bulgaria. By V. Beshkov 
and K. Nanev. 2006. Pensoft Publishers, Geo Milev Street 
13a 1111 Sofia, Bulgaria. 120 pages, Eur. 34.00 


* Antipredator Defenses in Birds and Mammals. By T. 
Caro. 2005. The University of Chicago Press 1427 East 60" 
Street, Chicago, Illinois 60637 USA. 592 pages, U.S. $38 


Retracing the Aurochs: History, Morphology and Ecolo- 
gy of an Extinct Wild Ox. By C. Van Vuure. 2005. Pensoft 
Publishers, Geo Milev Street 13a 1111 Sofia, Bulgaria. 424 
pages, Eur. 54.90 


The Return of the Beaver. By G. Sjoberg and J. Ball. 2006. 
Pensoft Publishers, Geo Milev Street 13a 1111 Sofia, Bulgaria. 
300 pages, Eur. 67.90 


A Systematic List of Extant Ground Beetles of the World 
(Coleoptera ‘‘“Geadephaga”’: Trachypachidae and Cara- 
bidae, Incl. Paussinae, Cicindelinae, Rhysodinae). Second 
Edition. By W. Lorenz. 2005. Pensoft Publishers, Geo Milev 
Street 13a 1111 Sofia, Bulgaria. 530 pages, Eur. 135.00 


To See Every Bird on Earth: a Father, a Son and a Life- 
long Obsession. By Dan Koeppel. 2005. Hudson Street Press, 
published by Penguin Group. 304 pages, U.S.$24.95 Cloth. 


* Rare Bird: Pursuing the Mystery of the Marbled Mur- 
relet. By Maria Mudd Ruth. 2005. Rodale Press, 33 East 
Minor Street, Emmaus, Pennsylvania 18098-0099. 298 pages, 
US. $23:95'Cloth. 


Birds of South Asia. The Ripley Guide. 2 volumes. By 
Pamela C. Rasmussen and John C. Anderton. 2005. Lynx 
Edicions, Montseny, 8, E-08193 Bellaterra, Barcelona, Spain. 
£55.00 Cloth 


Cercopid Spittle Bugs of the New World (Hemiptera, 
Auchenorrhyncha, Cercopidae). By C. Carvalho and M. 
Webb. November, 2005. Pensoft Publishers, Geo Milev Street 
13a 1111 Sofia, Bulgaria. 280 pages, Eur. 58.59. 


Curassows and Related Birds. By Jean Delacour and Dean 
Amadon. (Original 1973. Revised by D. Brooks). 2005. 
Lynx Edicions, Montseny, 8, E-08193 Bellaterra, Barcelona, 
Spain. 476 pages, U.S. $75. 


Ephemeroptera of South America. Aquatic Biodiversity 
of Latin America. (Abla Series) Number 2. By E. Domin- 
guez, C. Molineri, M. Pescador, M. Hubbard, C. Nieto. 2006. 
Pensoft Publishers, Geo Milev Street 13a 1111 Sofia, Bulgaria. 
490 pages, Eur. 95.00. 


BOOK REVIEWS 


47] 


ing pure denial of facts presented in this book, sug- 
gesting another period of “intellectual stagnation”. As 
a reviewer, I recommend all managers buy this book 
and implement its lessons learnt today. 

FALK HUETTMANN 


Institute of Arctic Biology, Biology and Wildlife Department, 
University of Alaska, Fairbanks Alaska 99775 USA 


Fishes of the River Cai, Vietnam, Khanh Hoa Province. 
By D. Serov. 2005. Pensoft Publishers, Geo Milev Street 13a 
1111 Sofia, Bulgaria. 164 pages, Eur. 33.00 


Coastal Fishes of Southern Africa. By Phil Heemstra and 
Elaine Heemstra. 2005. National Inquiry Services Centre. 
Distributed by NHBS2-3 Wills Road, Totnes, Devon TQ9 
5XN, UK 488 pages. £25.00 approximately U.S.$46 


The Frogs of New Guinea and the Solomon Islands. By J. 
Menzies. 2006. Pensoft Publishers, Geo Milev Street 13a 
1111 Sofia, Bulgaria. 210 pages. Eur 45.00 


Evolution of the Insects. By David Grimaldi and Michael 
S. Engel. 2005. Cambridge University Press, The Edinburgh 
Building, Shaftesbury Road, Cambridge, CB2 2RU UK. 
Distributed by NHBS2-3 Wills Road, Totnes, Devon TQ9 5XN, 
UK £45.00 Cloth 


Fascinating Insects. Some Aspects of Insect Life. By P. 
Jolivet and K.Verma. October, 2005. Pensoft Publishers, Geo 
Miley Street 13a 1111 Sofia, Bulgaria. 320 pages. Eur. 34.95 


Great British Marine Animals. By Paul Naylor. 2005 . 
Sound Diving Publications, Distributed by NHBS 2-3 Wills 
Road, Totnes, Devon TQ9 5XN, UK. 272 pages. £15.00 paper. 


An Atlas of the Reptiles of North Eurasia. By N. Ananjeva, 
N. Orlov, R. Khalikov, I. Darevsky, I. Ryabov and A. Bara- 
banov, 2006. Pensoft Publishers, Geo Milev Street 13a 1111 
Sofia, Bulgaria. 250 pages. Eur. 75.00 


_ Botany 


Illustrations of Alien Plants of the British Isles. By E. 
Clement, D. Smith and I. Thirlwell. 2005. Botanical Society 
of the British Isles, Botany Department, The Natural History 
Museum, Cromwell Road, London, SW7 5BD. 466 pages. 
£18.75 Paper, approximately U.S. $35 


* Plant Conservation — A Natural History Approach. 
Edited by G. Krupnick and W. Kress. 2005. The University 
of Chicago Press 1427 East 60" Street, Chicago, Illinois 
60637 USA. 344 pages. U.S.$30 


Prairie Phoenix: The Red Lily in Saskatchewan. Edited 
by Bonnie J. Lawrence and Anna L. Leighton. 2005. Nature 
Saskatchewan, Room 206, 1860 Lorne Street, Regina, Sas- 
katchewan S4P 2L7. 139 pages. $24.95 Cloth. 


* Giant Trees of Western North America and the World. 
By A. Carder. 2005. Harbour Publishing, Box 219 Madeira 
Park, British Columbia VON 2HO0. 152 pages. $26.95 Paper. 


* Tropical Rainforests — past Present and Future. By E. 
Bermingham. Edited by C. Dick and C. Moritz. 2005. The 
University of Chicago Press, 1427 East 60" Street, Chicago, 
Illinois 60637 USA. 672 pages. U.S. $45 Paper 


472 


Environment 


Crooked Lake Biophysical Survey 2000. By C. and Deirdre 
Griffiths. 2005. Alberta Environmental Network, 1-6328A 
104 Street NW, Edmonton, Alberta T6H 2K9. 330 pages. $90 
(Postage and handling, add $10) 


* Ecology & Evolution in the Tropics: A Herpetological 
Perspective. Edited by Maureen A. Donnelly, Brian I. Crother, 
Craig Guyer, Marvalee H. Wake, and Mary E. White. 2005. 
The University of Chicago Press, 1427 East 60" Street, Chicago, 
Illinois 60637 USA. 675 pages. U.S. $45. 


Scientists Debate Gaia. Edited by S. Schneider, J. Miller, E. 
Crist, and P. Boston. 2005. MIT Press, Five Cambridge Cen- 
ter, 4° Floor, Cambridge, Massachusetts 02142-1493. 400 
pages, US $ 50 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


An Ocean Odyssey. By Stephen Wong and Takako Uno. 
2005. Distributed by NHBS2-3 Wills Road, Totnes, Devon 
TQ9 5XN, UK. 240 pages, £30.00 Cloth 


One Planet, Many People: Atlas of Our Changing Envi- 
ronment. UNEP. Distributed by NHBS2-3 Wills Road, Totnes, 
Devon TQ9 5XN, UK. 320 pages, Hardcover £81.95 or down- 
load at http://www.na.unep.net/OnePlanetManyPeople/Atlas 
Download.php 


Tiger Bone and Rhino Horn — The Destruction of Wildlife 
for Traditional Chinese Medicine. By Richard Ellis. 2005. 
Distributed by NHBS2-3 Wills Road, Totnes, Devon TQ9 
S5XN, UK £16.95 paper 


* Sods, Soil, and Spades: Acadians at Grand Pre and their 
Dykeland Legacy. By J. Sherman Bleakney. 2004. McGill- 
Queen’s University Press, Montreal, Quebec. 221 pages, Can 
$106.99 


News and Comment 


The Boreal Dip Net/L'Epuisette Boreale: Newsletter of the Canadian Amphibian and Reptile 
Conservation Network/ Réseau Canadien de Conservation des Amphibiens et des Reptiles 


9(3) July 2005 


Editor’s Note (Kerrie Serben) — Rounding up the facts 
on Roundup R (Bruce Pauli and Christina Howe — Queries 
to CARCNET — Life on the Info Line (David Galbraith) — 
Baby it’s cold outside: Overwinter survival of Painted Turtle 
hatchlings (Elinor J. Hughes) — Interesting notes from Larry 
(Larry Halverson: Best management practices for amphib- 
ians and reptiles in urban and rural environments in BC; BC 
reptiles web site) — Blanding’s Turtles are Endangered in 
Nova Scotia — Will frogs derail the Sea to Sky Highway? — 
Guidance for determining riparian zones (Christine Bishop) — 


Marine Turtle Newsletter (109) 


July 2005. 28 pages: EDITORIAL: Tsunami Fund Update — 
GUEST EDITORIAL: An Economist’s Reflections on the 25" 
Annual Symposium for Sea Turtle Biology and Conservation: 
Empirical Program Evaluation and Direct Payments for Sea 
Turtle Conservation — Norges: Report of Olive Ridley Nesting 
on the North Coast of Trinidad — New Leatherback Conser- 
vation Project in Papua, Indonesia — A Note on the Illegal 
Trade in Stuffed Turtles in South Java, Indonesia — MEETING 
REPORTS — MTSG UPDATE — ANNOUNCEMENTS — OBITUARY — 
NEWS & LEGAL BRIEFS — RECENT PUBLICATIONS. 


Snake By D. H. Lawrence 1885-1930 (Sara Ashpole) — 10" 
Annual Meeting of CARCNET/RECCAR: Ottawa 15-19 
September 2005. 

— Membership in CARCNET/RECCAR is $20.00 (non- 
student) and $10 (student), and includes The Boreal Dipnet. 
Fees are due at the Annual Meeting (contact Bruce Pauli, Cana- 
dian Wildlife Service, National Wildlife Research Centre, 
Carleton University, Raven Road, Ottawa, Ontario KIA OH3. 
Web site: http://www.carcnet.ca/). 


The Marine Turtle Newsletter is edited by Brendan J. 
Godley and Annette C. Broderick, Marine Turtle Research 
Group, Centre for Ecology and Conservation, University of 
Exeter in Cornwall, Tremough Campus, Penryn TR10 9EZ 
United Kingdom; e-mail MTN @seaturtle.org; Fax +44 1392 
263700. Subscriptions and donations towards the production 
of the MTN can be made online at <http://www.seaturtle.org/ 
mtn/> or postal mail to Michael Coyne (online Editor) Marine 
Turtle Newsletter, | Southampton Place, Durham, North 
Carolina 27705 USA (e-mail: mcoyne @seaturtle.org). 


473 


Minutes of the 126 Annual Business Meeting of 
The Ottawa Field-Naturalists’ Club 11 January 2005 


Place and time: Canadian Museum of Nature, Ottawa, Ontario, 7:30 p.m. 


Chairperson: 
Attendance: 


Mike Murphy, President 


Twenty-eight persons attended the meeting. 


Attendees spent the first half-hour reviewing the minutes of the previous meeting, the Treasurer’s Report and 
the Report of Council. The meeting was called to order at 7:40 p.m. with some opening remarks from Michael 


Murphy, the President. 


1. Minutes of the Previous Meeting 

There were no changes to the minutes of the 125" 
Annual Business Meeting. 

It was moved by Fenja Brodo/Ron Bedford that the 
minutes be accepted. 

(Motion Carried) 


2. Business Arising from the Minutes 
There was no business arising from the Minutes. 


3. Communications Relating to the Annual 
Business Meeting 
There were no communications relating to the Annual 
Business Meeting. 


4. Treasurer’s Report 

Frank Pope reviewed the financial report for the year 
ending 30 September 2004, noting that the Club’s net 
assets have increased by approximately $18 000. Most 
of this came in dedicated funds. 

Moved by Frank Pope and seconded by Gillian 
Marston that the Financial Report be accepted. 

(Motion Carried) 


5. Committee Reports 

Mike Murphy introduced each of the Committee 
reports and a representative of the appropriate Com- 
mittee. After each report was read aloud, he asked for 
questions and comments. He thanked the committee 
chairs and committee members for their work over the 
past year. 

Moved by David Hobden/Otto Loesel, that the re- 
ports be accepted with suggested ammendments. 

(Motion Carried) 


6. Nomination of the Auditor 
Moved by Frank Pope, seconded by Dr. I. Brodo, 
that Janet Gehr continue as Auditor for another year. 
(Motion Carried) 


7. Report of the Nominating Committee 

Fenja Brodo expressed thanks to Michael Murphy 
(President), Gillian Marston (Vice-President) and the 
rest of Council on behalf of the club. Special thanks 
were extended to Eleanor Zurbrigg. 


President 

Vice President 
Secretary 
Treasurer 

Past President 
Business Manager 
Editor CFN 
Editor T&L 


Committee Chairs 
Birds 

Computers 
Conservation 
E&P 
E&L/Nominations 
Finance 

FWG 
Membership 
Publications 

FON Rep 


Members at large 
Kathy Conlan 
Diane Lepage 
Diane Kitching 
Henry Steger 


Chairs not on Council 


Awards 
Macoun 


Mike Murphy 


Susan Laurie Bourque 
Frank Pope 

Gary McNulty 

Bill Cody 

Francis Cook 

Karen McLachlan Hamilton 


Chris Traynor 
Eleanor Zurbrigg 
Stan Rosenbaum 
Gillian Marston 
Fenja Brodo 
Louise Schwartz 
David Hobden 
Dave Smythe 
Ron Bedford 
Cendrine Huemer 


Ernie Brodo 
Rob Lee 


Retiring from the council: John Cameron, 


New on the council: 


Barbara Gaertner, 
Christina Lewis 


Eleanor Zurbrigg 
(returning), 
Diane Kitching 


Moved by Fenja Brodo, seconded by Frank Pope, 
that the slate of nominations for the 2005 Council be 


accepted. 


(Motion Carried) 


8. New Business 


There was no new business. 


474 


2005 


9. Presentation by Robert Lee, ‘Mary Stuart 
and the Macoun Club” 

Rob Lee presented an enjoyable and touching pre- 
sentation documenting the long and fruitful association 
between the late Mary Stuart and members of the 
Macoun Club. Mary owned a large property in the area 
of Pakenham which she generously allowed the Macoun 
Club to use as a destination for both field and camping 
trips. Rob showed many slides taken over the past thirty 


MINUTES OF THE 126" ANNUAL BUSINESS MEETING 


475 


years, of her land and of Macoun members enjoying 
their time there. The slide show and Robs’ narration 
were very well received by club members. 


10. Adjournment 
Moved by Henry Steger/Ron Bedford that the meet- 
ing be adjourned at 9:35 pm. 
(Motion Carried) 


SUSAN LAURIE-BOURQUE, Recording Secretary 


The Ottawa Field-Naturalists’ Club Committee Reports for 2004 


Awards Committee 

The Awards Committee met in January to consider nomi- 
nations made for the various OFNC Awards. During the fol- 
lowing weeks and months, numerous communications were 
held by e-mail and Canada Post as well as by phone. As a 
result of our deliberations, the following five awards for the 
year 2003 were presented at the OFNC’s Annual Soirée, which 
took place 24 April 2004 at St. Basil’s Church in Ottawa. 


HONORARY MEMBER: 

Daniel Strickland — for his renowned studies on Gray Jays, 
and for the major role that he has played in fostering an ap- 
preciation and enjoyment of natural history by the general 
public. 


MEMBER OF THE YEAR: 

Pearl Peterkin for her hard work on the T&L back-issue 
project as well as all her involvement with the Education 
and Publicity Committee. 


GEORGE MCGEE SERVICE AWARD: 
Colin Bowen for his extraordinary work on the Birds Com- 
mittee Over many years. 


CONSERVATION AWARD — MEMBER: 

Daniel Brunton for his role in the establishment of a new 
Canadian chapter of the RiverKeeper Alliance in Ottawa, and 
securing a full-time RiverKeeper to monitor the health and 
conservation of the Ottawa River. 


CONSERVATION AWARD — NON-MEMBER: 

Friends of the Jock River for their dedicated, inspired, hard- 
working and thoughtful approach to protecting the health of 
the Jock River and its watershed. 

The full text of the citations for each of these awards was 
published in a previous issue of The Canadian Field-Naturalist. 

I. BRODO 


Birds Committee 

The Birds Committee participated together with the Club 
des Ornithologues de |’Outaouais in another successful Christ- 
mas Bird Count in December 2003. The Fall Bird Count was 
held in late October 2004 and proved continuingly popular. 
The Committee was again active with the Peregrine Falcon 
Watch at the downtown nest site. For the first time, an infor- 
mation sign was displayed and a donations box was mounted. 
Over $200 was collected. Both falcon chicks fledged suc- 
cessfully tin 2004. The Bird Record Sub-committee met dur- 
ing the year to review records of rare birds for the area. Re- 
sults of this work appear in Trail & Landscape. We continue 
to provide seasonal bird summaries for Trail & Landscap. 
The fourth year of surveying for the Ontario Breeding Bird 
Atlas (Region 24) has been completed. In 2004 the seed-a- 
thon raised over $600 in pledges for the OFNC’s bird feeders. 


We continue to operate the rare bird alert and Ottawa’s bird 
status line, which provides a recorded telephone message of 
current bird sightings. A new touch-tone feature added this 
year brings you directly to the most recent rare bird alert. 

C. TRAYNOR 


Computer Management Committee 

The Computer Management Committee (CMC) has been 
without a Chairman during 2004 and the Committee has not 
met during the year. In the absence of a Chairman, the 
President has acted on behalf of the Chairman at Council 
for any matters that would normally be dealt with by the 
Committee. 

In the almost 20 years since the CMC was first formed, 
there have been fundamental changes to computer technology 
such as the astonishing growth of the Internet and widespread 
use of personal computers by the general public. 

Despite this, the OFNC continues to rely on the same suite 
of computer applications, such as the OFNC membership 
database and the desktop publishing systems used for the 
Club publication (Trail & Landscape). That so little change 
has been needed shows how well-founded these applications 
were. Due to its complexity and the variety of vital infor- 
mation products it must provide, the Membership Database 
is the application most needing renewal. Work has com- 
menced on a replacement system. 

It should be noted that many of the newer computer-related 
services used by the Club, such as the Club Website at (http:// 
www.ofnc.ca) and e-mail addresses, have been established by 
Club volunteers without requiring direct involvement of the 
Computer Management Committee. 

M. MurRPHY 


Conservation Committee 
Larose Forest Francoscenie Proposal and Ontario Municipal 
Board Hearing 

Stan Rosenbaum appeared as a participant witness to sup- 
port the Friends of Larose Forest. OFNC approved payment 
of costs up to $1,000 to Federation of Ontario Naturalists for 
expert forestry witness Jim Faught, called by the Friends of 
Larose. 


South Gloucester Lands (5309 Bank Street) 

David Hobden represented OFNC at a City meeting. 
Numerous letters were sent to Ontario Ministry of Natural 
Resources and City Council, opposing quarrying and criti- 
cizing inadequate proposals to relocate rare species. 

Alfred Bog 
Frank Pope, chair of the Alfred Bog Committee, announced 


that the OMB appeal is abandoned and it now appears that 
Alfred Bog is protected. 


476 


Provincial Legislation Review 

Workshops attended on the Planning Act, Provincial Policy 
Statement (PPS), and Ontario Municipal Board (OMB). 
Written briefs and letters sent to the Ontario Ministries of 
Municipal Affairs, Environment, Natural Resources and 
Agriculture, and the Premier, and to an EA Advisory Panel. 


Constance Creek Wetland Donation Proposal 
The owner has offered this 50-acre property to the FON. 
A site visit was made by Frank Pope and Stephen Derbyshire 
of OFNC, and Chris Grooms of FON. 
S. ROSENBAUM 


Education and Publicity Committee 

The committee organized and managed OFNC participa- 
tion in four public events: the Ottawa Teachers Federation 
elementary teachers’ professional development day, 13 Feb- 
ruary 2004, the Carlingwood Wildlife Festival, 2 to 4 April; 
Telesat Canada’s Health Fair (theme Go Green) 12 May; 
Health Canada’s Celebration of Environment Week, 4 June. 
Arrangements were made for the judging and presentation 
of OFNC Natural History Awards at the Ottawa Regional 
Science Fair, 3 April. A nature photography contest for mem- 
bers was developed and details announced on the web-site 
and in volume 38 number 4 Trail & Landscape. 

A portable display was developed and first used on a fall 
weekend in Mer Bleue. A French edition of the club brochure 
was published and distributed. 

The sales table at the club’s evening presentation at the 
Canadian Museum of Nature provided revenue of $528. 

J. CAMERON 


Excursions and Lectures Committee 

In 2004 this Committee arranged 33 events, ten monthly 
meetings (including the Annual Business Meeting) and the 
Soirée. Of the 33 events, ten were general interest excursions, 
eight focused on birds, five on plants, five on invertebrates 
(butterflies, odonates, mollusca), two workshops (GPS and 
sketching) and four were museum trips. We hired buses for 
two of our trips: Presquile and the Biod6me in Montreal. Full 
day trips to interesting places remain the most popular and 
this year these included a walk in the Larose Forest (18 par- 
ticipants) and hiking up Luskville Falls (23 participants). 
Signed waiver forms allow us to track popularity of events. 

Topics covered in our monthly meetings were insects (two 
talks), plants as de-contaminators, Australia, Petrie Island, 
canoeing down the Saskatchewan River, Macoun Marsh 
Project and the popular members slide night. An amusing 
video on penguins filled in for a cancelled speaker. 

Council approved a budget of $200 for gifts/ honoraria to 
our speakers; the first was presented to our speakers in No- 
vember 2004. 

F. BRODO 


Executive Committee 
The Executive Committee met twice during 2004. The 
Committee discussed strategic planning matters such as gov- 
ernance, succession planning, and measures to address mem- 
bership decline within the Ottawa Field-Naturalists’ Club 
and the Macoun Field Club. Plans are also being developed to 
coordinate and acknowledge the efforts of volunteers, espe- 
cially those who are new members. 
M. MurpHy 


Finance Committee 
The Finance Committee met three times in 2003-04. High- 
lights include: 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


1. The Committee recommended to Council that no changes 
be made to the membership fee structure for 2005. Council 
approved this recommendation. Some concerns were noted 
about gradually declining membership and related revenue, 
operating deficits experienced over the past few years, and 
the reductions to the unrestricted reserve to cover these 
deficits. 

2. A proposed budget for 2004-05 for the OFNC and the 
CEN was submitted to Council. A deficit of over $17 000 is 
forecast, and if realized will be a draw on the unrestricted 
reserve. Council approved this proposed budget. 

During the year the Committee discussed the impact of 
changes in the federal government’s Publication Assistance 
Program on mailing costs for The Canadian Field-Naturalist 
and Trail and Landscape, donations that qualify within our 
Registered Charity status, and governance issues associated 
with club liability. 

L. SCHWARTZ 


Fletcher Wildlife Garden 

The year began with the signing of the first direct agreement 
between OFNC and Agriculture and Agri-food Canada con- 
cerning the operation of the Fletcher Wildlife Garden (FWG). 

For the OFNC Annual Business Meeting, in January 2004, 
we made a digital presentation on the history and activities 
of FWG, which combined slides, prints and digital pictures. 
We had displays at the Ottawa Home Show and the Wildlife 
Festival and at a Festival workshop at the Museum of Nature. 
We staged our own International Migratory Bird event in 
conjunction with the Festival. 

When the new gardening season began in April, all volun- 
teers were provided with written safety instructions and asked 
to sign a waiver form. Volunteers contribute about 2500 hours 
each year. This summer three groups operated at different 
times. 

In June, our annual plant sale included native plants grown 
by volunteers and by the Algonquin College Horticulture 
Program. It raised over $2000. 

This year we were able to employ a full-time summer stu- 
dent, using a grant from Human Resources and Skills Devel- 
opment Canada. In addition to working with visitors, she re- 
organized the library and worked on invasive plant control. 

Invasive plant control has concentrated on eliminating or 
reducing seed production by buckthorn and swallow-wort. 
We were warned that biocontrol would be the only effective 
long term control for swallow-wort, but it was at least ten 
years away. We have begun our own experiments on how 
best to contain the swallow-wort in the meantime. We have 
also used the web site and email to warn local government 
and community agencies of the problem. 

Frank Pope completed two terms as FWG/OFNC represen- 
tative on the Central Experimental Farm Advisory Committee 
and was replaced by Henry Steger. 

D. HOBDEN 


Macoun Field Club Committee 

The Committee met once during the year and otherwise 
communicated regularly to plan the program for the children 
and young people during the year. Committee members super- 
vised or gave presentations at 44 indoor meetings and led 
15 field trips, as well as 2 camping trips. 

The Club’s annual publication, The Little Bear, was pro- 
duced on time by one of the Senior members, without super- 
vision, but the newsletter had to be taken over by one of the 
Committee members. Membership in the Senior group was 
faltering. 


2005 
CANADIAN 

Type Local Other 
Family 301 (317) 19 (23) 
Individual 310 (323) VO CLL) 
Honorary 14 (15) 10 (9) 
Life 21 (21) 20 (20) 
Sustaining 1] (11) 4 (3) 
Total 657 (687) 160 (166) 


Extensive renovations at the Victoria Memorial Museum 
Building required the Club to vacate the room where our col- 
lections and library had been housed for the past 20 years. 
Almost all the equipment and collections were put into stor- 
age with the Canadian Museum of Nature, with only the most 
essential materials being kept available in locked cabinets in 
the Museum’s activity room. The entire library (1500 vol- 
umes) was moved to the home of one Committee member, 
and linked to a subject index on the Club’s website, so that 
books remain available to members. The renovations are 
expected to last several years. 

The Club’s website continues to be developed by the 
Committee members, with some content (meeting and field- 
trip reports) being written by members. 

R. LEE 


Membership Committee 
The distribution of memberships for 2004 is shown in the 
table (above), with the comparable numbers for 2003 in 
brackets. These statistics do not include the 23 affiliate orga- 
nizations which receive free copies of the Club’s publications. 
This year, the Club lost a long time member and strong 
supporter of the Macoun Field Club with the death of Mary 
Stuart. Mary joined the Club in 1944 and was later given an 
Honorary membership for her contributions to the Club. 
D. SMYTHE 


MINUTES OF THE 126" ANNUAL BUSINESS MEETING 


477 


FOREIGN 
USA Other Total 
| (1) l (1) 322 (342) 
io (24) 4 (6) 440 (464) 
0 (0) 0 (0) 24 = (24) 
6 (7) l (1) 48 (49) 
0 (O) 0 (O) 1d-. (14) 
26 (32) 6 (8) 849 (893) 


Publications Committee 

The Publications Committee met 3 times in 2004. 

Four issues of The Canadian Field-Naturalist were pub- 
lished in 2004: Volume 117, #2, 3, 4 and Volume 118, #1. 
These four issues contained 702 pages; 59 articles; 28 notes; 
95 book reviews; 115 new titles; 1 commemorative tribute; 
20 pages of News and Comments; | page of miscellany; and 
a 30 page index. To commemorate the 125" anniversary of The 
OFNC, a 40 page history of The Club authored by Dan Brunton 
appears in Volume 118, #1. Only one article qualified for sup- 
port from the Manning Memorial Fund for a total of $1440. 
The CFN is still running behind schedule; nevertheless a 
substantially higher number than in 2003 of articles, notes, 
and book reviews were published. 

Volume 38 of Trail & Landscape was published in four 
issues containing 200 pages. Special red covers commem- 
orated The Club’s 125" anniversary. Greatest emphasis was 
on birds, the Fletcher Wildlife Garden, and Club affairs. 

Publishing costs continue to increase, giving rise to finan- 
cial concern for The Club. 

R. BEDFORD 


478 


Auditor’s Report 


THE CANADIAN FIELD-NATURALIST 


To The Members of THE OTTAWA FIELD NATURALISTS’ CLUB 


I have audited the balance sheet of THE OTTAWA 
FIELD-NATURALISTS’ CLUB as at September 30, 2004, 
the statement of changes in net assets, and the state- 
ments of operations. These financial statements are 
the responsibility of the organization’s management. 
My responsibility is to express an opinion on these 
statements based on my audit. 

Except as explained in the following paragraph, I 
conducted my audit in accordance with Canadian gen- 
erally accepted auditing standards. Those standards 
require that I plan and perform an audit to obtain rea- 
sonable assurance whether the financial statements are 
free of material misstatement. An audit includes exam- 
ining evidence supporting the amounts and disclosures 
in the financial statements. An audit also includes as- 
sessing the accounting principles used and significant 
estimates made by management, as well as evaluating 
the overall financial statement presentation. 

In common with many non-profit organizations, the 
Ottawa Field-Naturalists’ Club derives some of its rev- 
enue from donations and fund raising activities. These 
revenues are not readily susceptible to complete audit 
verification, and accordingly, my verification was lim- 
ited to accounting for the amounts reflected in the 
records of the organization. 

In my opinion, except for the effect of the adjust- 
ments, if any, which I might have determined to be 
necessary had I been able to satisfy myself concerning 
the completeness of the revenues referred to in the 
preceding paragraph, these financial statements present 
fairly, in all material respects, the financial position of 
the OFNC as at September 30, 2004, and the results 
of its operations and changes in net assets for the year 
then ended in accordance with Canadian generally 
accepted accounting principles. 


JANET M. GEHR 
Chartered Accountant 
North Gower, Ontario 
3 January 2005 


Vol. 119 
The Ottawa Field-Naturalists’ Club 
Balance Sheet 
September 30, 2004 
2004 2003 
ASSETS 
CURRENT 
Cash (Note 1) $ 22,306 $ 2,781 
Investment certificates (Note 1) 29,175 43,764 
Marketable securities (Note 2) 261,050 246,467 
Accounts receivable 16,703 27,690 
Prepaid expenses 1,000 1,000 
330,234 321,702 
CAPITAL ASSETS (Note 3) _ - 
LAND — ALFRED BOG 3,348 3,348 
333,582 $325,050 
LIABILITIES AND FUND BALANCES 
CURRENT 
Accounts payable and 
accrued liabilities $ 2,000 $ 2,000 
Deferred revenue 11,714 122075 
SIF | eee 
LIFE MEMBERSHIPS 13,607 12,663 
NET ASSETS 
Unrestricted 62,667 60,609 
Club reserve 100,000 100,000 
Manning principal 100,000 100,000 
Manning interest - OFNC 2,457 1,434 
- CFN 15,805 13,153 
Seedathon Le2Sit 1,618 
Anne Hanes memorial 870 870 
de Kiriline-Lawrence 18,429 18,798 
Macoun Baillie Birdathon 1,052 [337 
Alfred Bog 3,700 493 
306,261 298,312 
333,582 $325,050 


2005 


The Ottawa Field-Naturalists’ Club 
Statement of Operations 
For the Year Ended September 30, 2004 


2004 2003 2004 2003 
REVENUE REVENUE 
Memberships $ 13,969 $ 14,615 Memberships $ 9,313 $ 9,743 
Trail and Landscape 280 196 Subscriptions 23,956 25,369 
Interest 1,835 1,339 Reprints 5,574 9,534 
GST rebate 768 780 Publication charges 23,685 50,559 
Fletcher Wildlife Garden (Note 4) 1,734 — Interest and exchange 8,290 7,940 
Other 373 617 GST rebate 3,889 3,040 
Tr co =e Other L477 734 
18,959 17,547 — ne 7 
eae _ 76,164 = 106,919 
OPERATING EXPENSES 
Affiliation fees 670 ead, | EABHye 
Computer 1.294 302 Publishing 46,385  $ 80,758 
Membership 1,168 1,151 Reprints 2,141 7,307 
Office Assistant 1,000 1,000 Circulation 6,442 11,307 
Telephone 1,640 1,622 Editing 3,587 4,183 
Insurance 725 655 Office Assistant 5,000 5,000 
Audit i 000 1 000 Honoraria 9,000 9,000 
Car 1,214 1,290 GST rebate 4,203 6,508 
Other 952 884 Other = ON) eg a 
9.663 8.584 76,823 124,615 
=a a EXCESS EXPENSES OVER 
CLus ACTIVITY EXPENSES REVENUE $ (659) $ (17,696) 
Awards 150 ie 
Birds 540 1,696 
Education and Publicity VIS 73 
Excursions and Lectures (508) 496 
Macoun Field Club 606 851 
Soiree 270 19 
Trail and Landscape hate’ 8,368 
Fletcher Wildlife Garden (Note 4) - 2,025 
Other 84 _ 
11,495 14,126 
2 ASS 22710 
EXCESS EXPENSES OVER 
REVENUE $ (2,199) $(5,163) 


MINUTES OF THE 126'™ ANNUAL BUSINESS MEETING 


The Ottawa Field-Naturalists’ Club 


479 


Canadian Field-Naturalist — Statement of Operations 


For the Year Ended September 30, 2004 


480 THE CANADIAN FIELD-NATURALIST 


The Ottawa Field-Naturalists’ Club Notes to the Financial Statements 
September 30, 2004 


1. CASH 


Chequing 

Savings 

Nesbitt Burns 

Fletcher Wildlife Garden 


Investment Certificates: Maturity Maturity 
Value Date 
$ 29,402 04/29/05 


2. MARKETABLE SECURITIES 


Investment Certificates: Maturity Maturity 
Value Date 
Province of Newfoundland Coupon $ 44,782 10/17/11 
Province of Ontario Coupon 153376 12/02/12 
Province of Manitoba Coupon 29,847 11/15/04 
Ontario Savings Bonds 40,000 06/21/05 
CMHC Global Debs 52,000 12/01/06 
Province of Ontario Bond 30,000 09/12/07 
Province of Newfoundland Bond 20,000 10/07/08 
Government of Canada Coupon 30,167 12/01/09 
Province of New Brunswick Bond 20,000 06/15/10 


3. CAPITAL ASSETS 
Equipment at a cost of $16,748 is fully amortized. 


4, FLETCHER WILDLIFE GARDEN 


REVENUE 

Human Resources and Skills Dev. Canada 
TD Friends of the Environment Fund 
Taverner Cup 

Sales 

GST 

Donations 

Other 


EXPENSES 

Program 

Backyard 

Habitats 
Interpretation centre 
Administration 
Publications 

GST 

Library 


2004 


$ 11,899 
4,491 


5.96 
$ 22,306 


Yield 


1.35% 


Yield 


4.525% 
4.591% 
5.119% 
6.45% 

5.250% 
6.125% 
6.263% 
5.605% 
6.231% 


Vol. 119 


2003 


S (3,792) 
1,344 

oe 
4,295 


$ 2,781 


Book 
Value 
$ 29,175 


Book 
Value 
$ 32,965 
10,601 
27,939 
41,370 
53,539 
31,187 
20,538 
21,843 
21,068 


$261,050 


243 
132 


6,346 
$ (2,623) 


2005 


MINUTES OF THE 126'™ ANNUAL BUSINESS MEETING 


48] 


The Ottawa Field-Naturalists’ Club Statement of Changes in Net Assets 


For the Year Ended September 30, 2004 (Note 5) 


Net Beginning Excess 
Assets Balance Expenses CEFN 
Unrestricted $ 60,609 $ (659) 
Club Reserve 100,000 - 
Manning Principal 100,000 — 
Manning - OFNC 1,434 - 
Manning - CEFN 13,153 - 
Seedathon 1,618 = 
Anne Hanes Memorial 870 - 
de Kiriline-Lawrence 18,798 _ 
Macoun Baillie Birdathon hes erg _ 
Alfred Bog 493 — 
$ 298,312 $ (659) 


5. STATEMENT OF CHANGES IN NET ASSETS 


Excess Other 2004 Ending 
Expenses OFNC Revenue Expenses Balance 
$ (2,199) $ 4,916 $ - §$ 62,667 
- - - 100,000 
— ~ ~ 100,000 
- 1025 — 2,457 
- 4,092 1,440 15,805 
— 623 960 1,281 
~ ~ _ 870 
- 631 1,000 18,429 
- 115 400 1,052 
- 3,207 - 3,700 
$ (2,199) $ 14,607 $ 3,800 $306,261 


a) Manning CFN Expenses: Volume 117, Number 3, Cody, Reading and Line, Additions and range extensions to the 
vascular plant flora of the continental NWT and Nunavut, Canada, II. 


b) Seedathon Expenses: birdseed for club feeders. 


c) Louise de Kiriline Lawrence Expenses: Contribution to the defense of the Larose Forest at a hearing of the Ontario 


Municipal Board. 


d) Macoun Baillie Birdathon: a digital camera with accessories. 


The Ottawa Field-Naturalists’ Club Summary of Significant Accounting Policies 


September 30, 2004 
1. Nature of Business 

The organization is non-profit and incorporated under the 
laws of Ontario (1884). The organization promotes the appre- 
ciation, preservation, and conservation of Canada's natural 
heritage. It encourages investigation and publishes the results 
of the research in all fields of natural history and diffuses in- 
formation on these fields as widely as possible. It also supports 
and cooperates with other organizations engaging in preserv- 
ing, Maintaining or restoring environments of high quality 
for living things. 


2. Financial Instruments 

The organization's financial instruments consist of cash, 
accounts receivable, marketable securities, and accounts pay- 
able. Unless otherwise noted, it is the management's opinion 
that the organization is not exposed to significant interest, 
currency, or credit risks arising from these financial instru- 
ments. The fair value of these instruments approximate their 
carrying values. 


3. Capital Assets 

Capital assets acquired after 1989 are expensed. Capital 
assets acquired prior to 1990 were recorded as assets at cost 
and amortized on a stright-line basis. These assets have been 
fully amortized. 


4. Statement of Changes in Financial Position 

A statement of changes in financial position has not been 
provided as it would not provide additional meaningful in- 
formation. 


5. Foreign Currency 

Transactions during the year in U.S. dollars have been con- 
verted in the accounts to Canadian dollars at the exchange 
rate effective at the date of the transaction. All monetary 
assets in U.S. dollars at year end have been converted to 
Canadian dollars at the rate effective on Sept. 30 2004. Gains 
or losses resulting therefrom are included in revenue or 
expenses. 


Advice for Contributors to The Canadian Field-Naturalist 


Content 

The Canadian Field-Naturalist is a medium for the publi- 
cation of scientific papers by amateur and professional natu- 
ralists or field-biologists reporting observations and results 
of investigations in any field of natural history provided that 
they are original, significant, and relevant to Canada. All read- 
ers and other potential contributors are invited to submit for 
consideration their manuscripts meeting these criteria. The 
journal also publishes natural history news and comment items 
if judged by the Editor to be of interest to readers and sub- 
scribers, and book reviews. Please correspond with the Book 
Review Editor concerning suitability of manuscripts for this 
section. For further information consult: A Publication Policy 
for the Ottawa Field-Naturalists’ Club, 1983. The Canadian 
Field-Naturalist 97(2): 231-234. Potential contributors who 
are neither members of The Ottawa Field-Naturalists’ Club 
nor subscribers to The Canadian Field-Naturalist are encour- 
aged to support the journal by becoming either members or 
subscribers. 


Manuscripts 

Please submit, to the Editor, in either English or French, 
three complete manuscripts written in the journal style. 
The research reported should be original. It is recommended 
that authors ask qualified persons to appraise the paper before 
it is submitted. All authors should have read and approved it. 
Institutional or contract approval for the publication of the data 
must have been obtained by the authors. Also authors are ex- 
pected to have complied with all pertinent legislation regard- 
ing the study, disturbance, or collection of animals, plants or 
minerals. The place where voucher specimens have been de- 
posited, and their catalogue numbers, should be given. Lati- 
tude and longitude should be included for all individual local- 
ities where collections or observations have been made. 

Print the manuscript on standard-size paper, doublespace 
throughout, leave generous margins to allow for copy mark- 
ing, and number each page. For Articles and Notes provide 
a citation strip, an abstract and a list of key words. Gener- 
ally, words should not be abbreviated but use SI symbols for 
units of measure. The names of authors of scientific names may 
be omitted except in taxonomic manuscripts or other papers 
involving nomenclatural problems. “Standard” common names 
(with initial letters capitalized) should be used at least once 
for all species of higher animals and plants; all should also 
be identified by scientific name. 

The names of journals in the Literature Cited should be 
written out in full. Unpublished reports and web documents 
should not be cited here but placed in the text or in a sepa- 
rate Documents Cited section. List the captions for figures 
numbered in arabic numerals and typed together on a separate 
page. Present the tables each titled, numbered consecutively 
in arabic numerals, and placed on a separate page. Mark in 
the margin of the text the places for the figures and tables. 


Check recent issues (particularly Literature Cited) for 
journal format. Either “British” or “American” spellings are 
acceptable in English but should be consistent within one 
manuscript. The Oxford English Dictionary, Webster’s 
New International Dictionary and le Grand Larousse 
Encyclopédique are the authorities for spelling. 


Illustrations 

Photographs should have a glossy finish and show sharp 
contrasts. Electronic versions should be high resolution. Photo- 
graphic reproduction of line drawings, no larger than a 
standard page, are preferable to large originals. Prepare line 
drawings with India ink on good quality paper and letter (don’t 
type) descriptive matter. Write author’s name, title of paper, and 
figure number on the lower left corner or on the back of each 
illustration. 


Reviewing Policy 

Manuscripts submitted to The Canadian Field-Naturalist 
are normally sent for evaluation to an Associate Editor (who 
reviews it or asks another qualified person to do so), and at 
least one other reviewer, who is a specialist in the field, cho- 
sen by the Editor. Authors are encouraged to suggest names 
of suitable referees. Reviewers are asked to give a general 
appraisal of the manuscript followed by specific comments 
and constructive recommendations. Almost all manuscripts 
accepted for publication have undergone revision—sometimes 
extensive revision and reappraisal. The Editor makes the 
final decision on whether a manuscript is acceptable for pub- 
lication, and in so doing aims to maintain the scientific quality, 
content, overall high standards and consistency of style, of 
the joumal. 


Special Charges — Please take note 

Authors must share in the cost of publication by paying 
$80 for each page, plus $15 for each illustration (any size up 
to a full page), and up to $80 per page for tables (depending 
on size). Authors may also be charged for their changes in 
proofs. Reproduction of colour photos is extremely expensive; 
price quotations may be obtained from the Business Manager. 
If grant or institutional funds are not available, club members 
and subscribers may apply for a waiver of charges for the 
first five pages. 

Limited joumal funds are available to help offset publi- 
cation charges to authors with minimal financial resources. 
Requests for financial assistance should be made to the Busi- 
ness Manager when the manuscript is accepted. 


Reprints 
An order form for the purchase of reprints will accompany 
the galley proofs sent to the authors. 


FRANCIS R. Cook, Editor 
R.R. 3 North Augusta, Ontario KOG IRO Canada 


482 


TABLE OF CONTENTS (concluded) Volume 119 Number 3 


Notes 
Possible tool use by Beavers, Castor canadensis, in a northern Ontario watershed D. M. BARNES 


Predation of a bat by American Crows, Corvus brachyrhyncos 
KARA L. LEFEVRE and CHRISTOPHER R. T. SULWAY 


Mortality of Deer Mice, Peromyscus maniculatus, in wire mesh live-traps: 
A cautionary note THOMAS S. JUNG and KIERAN S. O’ DONOVAN 


Longevity and productivity of three Wolves, Canis lupus, in the wild 
JIM HOLYAN, DIANE K. Boyb, Curtis M. MACK, and DANIEL H. PLETSCHER 


Observations of autumn courtship and breeding in Brown Bears, Ursus arctos, 
from coastal British Columbia OWEN T. NEVIN and BARRIE K. GILBERT 


Long-distance movement by a dispersing Deer Mouse, Peromyscus maniculatus, 
in the boreal forest THOMAS S. JUNG, KIERAN S. O’ DONOVAN, and TODD POWELL 


Movements of a two rabid Racoons, Procyon lotor, in eastern Ontario 
RICK ROSATTE, MIKE ALLAN, ROB WARREN, PETER NEAVE, TODD BABIN, LUKE BUCHANAN, 
DENNIS DONAVAN, KIRK SOBY, CHRIS DAVIES, FRANCES MULDOON, and ALEX WANDELER 


Common Loon, Gavia immer, nest attendance patterns recorded by remote video camera 
WING GOODALE, LEE ATTIX, and DAVID EVERS 


Short-tailed Shrew, Blarina brevicauda, apparently killed by Red Squirrel, 
Tamiasciurus hudsonicus ROBERT W. NERO 


Extension de |’aire de distribution de la Couleuvre a collier, Diadophis punctatus, 
dans l’est du Québec JEAN-FRANCOIS DESROCHES et BENOIT ROUSSEL 


Do juvenile Nearctic River Otters, Lontra canadensis, contribute to fall sent marking? 
ZACH H. OLSEN, SADIE S. STEVENS, and THOMAS I. SERFASS 


Book Reviews 
OOLOGY: Damselflies of Alberta: Flying Neon Toothpicks in the Grass 


OTANY: Flower Guide for Holiday Weekends — Orchids of Manitoba 


NVIRONMENT: Environmentalism Unbound: exploring New Pathways — The Love of Nature and the 
End of the World: The Unspoken Dimensions of Environmental Concern — Survival by Hunting: 
Prehistoric Predators and Animal Prey — Wildlife, Conservation and Human Welfare: A United States 
and Canadian Perspective 


NEw TITLES 


ews and Comment 

he Boreal Dip Net/L’Epuisette Boreale: Newsletter of the Canadian Amphibian and Reptile Conservation 
Network/ Reseau Canadien de Conservation des Amphibiens et des Reptiles 9(3) July 2005 — Marine 
Turtle Newsletter (109) July 2005 


Minutes of the 126th Annual Business Meeting of The Ottawa 
Field-Naturalists’ Club Tuesday 11 January 2005 


Advice to Contributors 


Mailing date of the previous issue 119(2): 13 June 2006 


2005 


44] 


443 


445 


447 


449 


451 


453 


455 


456 


457 


459 


462 
462 


463 
47] 


473 


474 


482 


THE CANADIAN FIELD-NATURALIST Volume 119 Number 3 


Articles 


An analysis of parasites of a mid-winter population of the Snowshoe Hare, Lepus americanus, 
on insular Newfoundland during a cyclical peak 
K. E. BENNETT, E. M. Bacaas, J. R. FINNEY-CRAWLEY, and M. MCGRATH 


Reconstructing changes in abundance White-tailed Deer, Odocoilus virginianus, Moose, 
Alces alces, and Beavers, Castor canadensis, in Algonquin Park, Ontario, 1860-2004 
NORMAN W. S. QUINN 


The distribution and habitat selection of introduced Grey Squirrels, 
Sciurus carolinensis, in British Columbia EMILY K. GONZALES 


The Flathead Minnow, Pimephales promelas, in New Brunswick 
DALE J. HooD and RUDOLPH F. STOCEK 


Vision and its relationship to novel behaviour in St. Lawrence Greenland Sharks, Somniosus 
microcephalus CHRIS J. HARVEY-CLARK, JEFFREY J. GALLANT, and JOHN H. BATT 


Characteristics d’une population introduit du Grand brochet, Esox lucius, dans 
le lac Ramsay, Parc de la Gatineau, Quebec, et impact sur l’ichthyofaune 
JOSIANE VACHON, BRIGITTE LAVALLEE, et FRANCOIS CHAPLEAU 


Woodpecker nest tree characteristics in upper midwestern oak forests 
COLLETTE L. ADKINS GIESE and FRANCESCA J. CUTHBERT 


Detectability of non-passerines using “pishing” in eastern Ontario woodlands 
J. RYAN ZIMMERLING 


Evidence of range expansion of eastern Coyotes, Canis latrans, in Labrador 
TONY E. CHUBBS and FRANK R. PHILLIPS 


Consequences of Beaver, Castor canadensis, flooding on a small shore fen 
in southwestern Quebec Joyce M. REDDOCH and ALLAN H. REDDOCH 


Long-eared Owls, Asio otus: A review of North American banding C. STUART HOUSTON 


Cyclopyxis acmodonta n.sp. and Arcella formosa n.sp.: two new species of testate rhizopods 
(Arcellinida, Protozoa) from threatened wetlands in Ontario, Canada 
KENNETH H. NICHOLLS 


A range extension for the Rock Vole, Microtus chrotorrhinus, in Labrador SARAH W. LANSING 


New records of vascular plants in the Yukon Territory VII 
WILLIAM J. CoDy, BRUCE BENNETT, and PHIL CASWELL 


Additions to the flora of the continental Northwest Territories from 
the Great Slave Lake area PAUL M. CATLING 


ISSN 0008-3550 


The CANADIAN 
FIELD-NATURALIST 


Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada 


Volume 119, Number 4 October—December 2005 


The Ottawa Field-Naturalists’ Club 


FOUNDED IN 1879 


Patrons 
Her Excellency The Right Honourable Michaélle Jean 
Governor General of Canada 


The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to 
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The Canadian Field-Naturalist 0.) <2, 


Volume 119, Number 4 October—December 2005 


Breeding and Non-Breeding Range of Canada, Branta canadensis, 
and Cackling Geese, Branta hutchinsii, in the Eastern Canadian Arctic 


MarkK L. MALLory!, ALAIN J. FONTAINE!, and HUGH Boypb? 


' Canadian Wildlife Service, P.O. Box 1714, Iqaluit, Nunavut XOA OHO Canada 

* Canadian Wildlife Service, National Wildlife Research Centre, Carleton University (Raven Road), Ottawa, Ontario K1A 0H3 
Canada 

3 Corresponding author: mark.mallory @ec.gc.ca; phone: (867) 975-4637 


Mallory, Mark L., Alain Fontaine, and Hugh Boyd. 2005. Breeding and non-breeding range of Canada, Branta canadensis, 
and Cackling Geese, Branta hutchinsii, in the eastern Canadian Arctic. Canadian Field-Naturalist 119(4): 483-489. 


The accepted breeding distribution of Canada Geese from the Atlantic Population (Branta canadensis interior) in the eastern 
Canadian Arctic is currently confined to northern Québec and the south coast of Baffin Island. Here we provide evidence 
based on observations from scientific studies, Inuit hunters, and territorial Wildlife Officers that B. c. interior now breeds in 
growing numbers 500 km farther north on northeastern Baffin Island than previously reported. Cackling Geese (B. hutchinsii), 
which breed more widely across eastern Arctic Canada, to about 72°N, may also be increasing there. Moreover, individuals 
of both species are seen occasionally as far north as Ellesmere Island in small flocks and within migrating or moulting 
flocks of Snow Geese (Chen caerulescens) or Brant (B. bernicla hrota), though none of these far northern stragglers are known 
to have bred. Whether these observations reflect a recent range expansion or improved distributional knowledge from more 


intensive recent survey efforts remains unknown. 


Key Words: Branta canadensis, Canada Goose, Branta hutchinsii, Cackling Goose, breeding, distribution, Arctic 


Our knowledge of the distribution of breeding birds 
in the Canadian Arctic is based largely on reports from 
early explorers (e.g., Parry 1824; Nansen 1897), from 
initial scientific survey efforts (e.g., Duvall and Han- 
dley 1948*; Ellis and Evans 1960; Manning 1976), 
and from more recent aerial reconnaissance (e.g., Net- 
tleship 1974; McLaren 1982; Gaston et al. 1986). Some 
long-term studies have also provided insights into 
annual variation in species that breed in certain areas 
(e.g., Gaston and Ouellet 1997; Lepage et al. 1998). 
While many gaps in our knowledge of breeding bird 
distributions in the Arctic remain, it is now clear that 
distributions of some birds in the Arctic are changing. 
For example, Snow Goose (Chen caerulescens) col- 
onies have expanded markedly in the past 30 years; 
breeding now occurs in new areas or at higher densi- 
ties than previously recorded, in some cases seriously 
degrading habitats (Mowbray et al. 2000). Ross’s 
Geese (Chen rossi) now breed farther east in Nunavut 
and in greater numbers than they did 20 years ago 
(Ryder and Alisauskas 1995). In addition, satellite 
telemetry has shown that some Canada Geese (Branta 
canadensis) use southeastern Baffin Island as a stop- 
Over point en route to their breeding areas in Green- 
land (Scribner et al. 2003). 


The breeding distribution of different populations 
of Canada Geese in the eastern Canadian Arctic has 
recently undergone extensive review (Dickson 2000a; 
Canadian Wildlife Service Waterfowl Committee 2003; 
Boyd and Dickson in Kear 2004). Even more recently, 
the American Ornithologists’ Union (Banks et al. 2004) 
decided to split Canada Geese into two species, sepa- 
rating the small Cackling Geese, B. hutchinsii, from 
the larger forms of B. canadensis. Two breeding pop- 
ulations of these types of similar geese can be com- 
monly found north of 60° in eastern Nunavut. In the 
Kivallig region and around Foxe Basin, the Tallgrass 
Prairie Population dominates, which is principally 
composed of Cackling Geese (Bellrose 1980; Dick- 
son 2000b). Some of the Atlantic Population geese 
(mostly B. c. interior, a larger race of Canada Goose) 
that breed in northern Québec may breed in southwest- 
ern Baffin Island. Although the North Atlantic Popu- 
lation (B. c. canadensis, also a larger race of Canada 
Goose) breeds in northern Labrador, it is not reported 
to breed on Baffin Island. To the east across Baffin 
Bay, Canada Geese breeding in western Greenland 
are morphologically and genetically similar to the 
Atlantic population (Fox et. al. 1996; Scribner et al. 
2003). 


483 


484 


We have combined breeding range data from a vari- 
ety of sources in Figure | to show the current, accepted, 
northern portion of the breeding distribution of Canada 
and Cackling geese (Bellrose 1980; Reed et al. 1980; 
Godfrey 1986; Dickson 2000b; Mowbray et al. 2002). 
Godfrey (1986) mentioned that some geese were 
thought to breed on the Cumberland Peninsula, with- 
out confirmation. This note deals chiefly with new in- 
formation on the distribution and probable range expan- 
sion of the Atlantic Population of B. c. interior into 
the eastern Canadian Arctic islands. It has also moved 
into west Greenland, where its numbers have grown 
rapidly in the last 30 years (Fox et al. 1996; Malecki 
et al. 2000; Scribner et al. 2003). This new information 
suggests that current range maps require revision. 


Methods 

Much of the information on the breeding range of 
Canada and Cackling geese reported here came from 
Inuit hunters, including verified data from the five-year 
Nunavut Wildlife Harvest Study (Priest and Usher 
2004). This harvest study required hunters to identify 
the number, location and date of animals (in this case, 
geese and/or eggs) harvested, and these records were 
subsequently verified by field technicians. These results 
were then summarized and published (Priest and Usher 
2004), such that results for a community represent 
harvest in that year within the hunting region around 
that area. Other data were also collected as part of local 
ecological knowledge studies focused on various sub- 
jects during discussions with Inuit hunters and Wildlife 
Resource Officers in communities along Baffin Bay 
(e.g., Mallory et al. 2003). Local knowledge has proven 
to be an effective means of examining wildlife distri- 
butions in the Arctic, as Inuit are keen observers of 
their environment (Gilchrist et al. 2005*). 

The other main sources of data were incidental 
observations by MLM and AJF in the course of ground- 
and boat-based surveys of other migratory bird species 
between 2000 and 2004 (Fontaine et al. 2001; Mallory 
et al. in review). Scattered published and unpublished 
materials collected by other biologists during non-sys- 
tematic aerial surveys of the Queen Elizabeth Islands 
and coastlines of Lancaster and Jones Sound in 1968- 
1969 and 1971 amplify some of those records (e.g., 
Heyland and Boyd 1969*; HB). In these cases, “large” 
or “small” geese were typically identified by the rela- 
tive size of their head and neck, or their body size in 
relation to nearby birds (e.g., Snow Geese). There have 
been no systematic surveys to determine the distribu- 
tion of Canada or Cackling geese in the eastern Cana- 
dian Arctic. However, portions of the breeding popu- 
lation have been assessed directly (e.g., Malecki and 
Trost 1990), as part of colony surveys for Snow Geese 
(D. Caswell, personal communication), or in some 
other bird survey projects (e.g., Johnston et al. 2000). 


THE CANADIAN FIELD-NATURALIST 


Vol 119 


Results 
Reports by Inuit hunters 

In discussions with hunters in eastern Nunavut, we 
received reports on Canada Geese near various com- 
munities. Hunters in Iqaluit (63°45'N 68°30'W) have 
observed increases in the number of large Canada 
Geese upon islands and hillsides of Frobisher Bay. In 
Pangnirtung (66°30'N 66°W), Inuit hunters and Nation- 
al Park wardens stated that numbers of geese in Cum- 
berland Sound have increased dramatically since the 
1970s, and that some birds breed in the area, often at 
eider colonies. Further north, hunters from Qikiqtar- 
juaq (67°30'N, 64°W) told us about harvesting adult 
geese and their eggs near Cape Searle (67°14'N, 
62°28'W) and Reid Bay (66°56'N, 61°46'W), the same 
area where we observed breeding geese during our field 
studies (below). At Clyde River (70°45'N, 68°W), 
local hunters observe and harvest medium to large- 
sized geese. In Arctic Bay (73°02'N, 85°10'W), hunters 
shoot adult small geese, presumably B. hutchinsii, but 
local breeding has not been confirmed, although Cack- 
ling Geese do breed near southern Admiralty Inlet (Fig- 
ure 1). 

Results from the Nunavut Wildlife Harvest Study 
support the local ecological knowledge shared with us 
by Inuit hunters. Between 1996 and 2001, adult geese 
(apparently large geese, hence Canada Geese) were 
harvested by hunters from the communities of Iqaluit, 
Pangnirtung, Qikiqtarjuagq, Clyde River, Pond Inlet, 
Resolute Bay, and Grise Fiord (Figure 1; Priest and 
Usher 2004). Harvest of Canada Geese was not report- 
ed at Grise Fiord between 1956 and 1972 (Riewe 
1977). The magnitude of the annual harvest decreas- 
es as one moves north from Iqaluit to Resolute, with 
approximately 350 Canada Geese harvested each year 
among these communities (Priest and Usher 2004). 
An estimated 350 Canada Goose eggs are collected an- 
nually among residents of Iqaluit, Pangnirtung, Qik- 
iqtarjuaq and Clyde River (Priest and Usher 2004), 
confirming breeding near these communities. 


Other observations of breeding Canada Geese 

During surveys of breeding marine birds along the 
coastline of Frobisher Bay and Cumberland Sound in 
August 2000, Canada Geese were seen frequently in 
lowland arctic meadows and on grassy hillsides facing 
the coast (Fontaine et al. 2001), many of them with 
young of the year (J. A. Akearok, personal communi- 
cation). In other breeding bird surveys along the north 
shore of Frobisher Bay in July 2001 and 2002, numer- 
ous other flocks of adults, as well as paired nesting 
Canada Geese were observed (MLM). In June 2000 
to 2004, numerous small flocks of paired birds with no 
young were observed near the Iqaluit airport and in 
nearby inland valleys. Hence, geese in the Frobisher 
Bay area appear to be a mix of breeding birds, failed 
breeders and moult migrants, all of medium to large 
races. 


2005 


425° 115° 105° 95° 85° 75° 65° 
200 400 Kilometers 


MALLorRY, FONTAINE, AND BOYD: CANADA AND CACKLING GEESE 


485 


Known range 

Canada Goose - confirmed breeding 
Canada Goose - non-breeding 
Canada Goose - harvest 

Cackling Goose - confirmed breeding 
Cackling Goose - non-breeding 

Not observed 


® 
we 
A 
© 
[=] 
* 


Davis 
Strait 


Qikigtarjuaq 


FIGURE |. Known range (shaded) and new location observations (dots) of Canada (Branta canadensis) and Cackling geese 
(Branta hutchinsii) in eastern Nunavut, Canada. The existing range represents a composite of information from a variety 
of published sources. Note that solid triangles for Canada Goose harvest include harvest of adults and/or eggs, and do 
not differentiate between Canada Geese and Cackling Geese. 


On 11-14 June 2001, we also observed approxi- 
mately 50 nesting Canada Geese on the northern 
Cumberland Peninsula near Merchants Bay (67°20'N, 
62°30'W). Inuit at a local outpost camp had been har- 
vesting Canada Goose eggs for a week; there were at 
least 12 eggs in a bucket at the camp on 11 June. Anoth- 
er 18 pairs of breeding geese were seen on 7 June 2002 
on nearby Qaqulluit Island (67°12'N, 62°33'W). Dur- 
ing the trips from Qikiqtarjuaq to Qaqulluit Island, 
other Canada Geese were often flushed from heath 
slopes and lowlands along the 100-km route. All the 
geese in this region were large, suggesting B. c. inte- 
rior affinities. 


On southern Bylot Island (73°N, 78°W), J. D. 
Heyland (personal communication) found nesting geese 
each year from 1969-1971. Most of them were small, 
apparently Cackling Geese, but in 1970 a larger pair 
bred successfully (JDH and HB). A photograph taken 
before 1937 at Pond Inlet shows a local Inuit woman 
with Snow Geese and one Cackling Goose harvested 
nearby (A. Reed, personal communication). Lepage 
et al. (1998) saw flocks of small, Cackling Geese on 
Bylot Island almost every year between 1979 and 1997, 
and confirmed breeding on three occasions. They also 
saw five larger birds that they suggested were B. c. in- 
terior. C. Machtans flushed a Canada Goose of un- 


486 


recorded size off a nest containing four eggs on 28 June 
1997 at Creswell Bay (72°50'N, 93°11"W) on Somerset 
Island (C. Machtans, unpublished data). Its mate was 
in the vicinity and other geese (approximately four) 
were also observed but details concerning breeding 
status were not recorded. 

Cackling or Canada geese were seen in apprecia- 
ble numbers along the east and west coasts of Foxe 
Basin in 1979 (Reed et al. 1980), but not on the islands 
in the Basin. In 1987 and 1988, small numbers of medi- 
um-sized geese bred on Rowley Island (69°N, 78°W; 
Boyd 1989). The parents of one brood were noticeably 
larger than those of a second brood. As none were 
caught and measured, it was not possible to determine 
the species involved. In 1989, at least five pairs of 
small geese (B. hutchinsii) were seen at nests in the 
northwest of Prince Charles Island (Boyd 1999). 


Adult Cackling and Canada geese in the Queen Eliza- 
beth Islands 

A flightless dark goose was seen among a flock of 
20 flightless Snow Geese at the north end of Vendom 
Fiord, Ellesmere Island (77°40'N, 82°30'W) on 26 July 
1968, and another in a flock of 70 Snow Geese on a 
small lake near Goose Fiord (76°57'N, 88°45'W), on 
8 August 1971 (HB). They were smaller than the Snow 
Geese, suggesting B. hutchinsii. No Canada Geese 
were seen on extensive aerial surveys of Devon and 
Ellesmere islands, nor along the coast of northwest- 
ern Greenland, in 1969. 

On 10 June 2003, a single, large race Canada Goose 
was photographed with a flock of Eastern High Arctic 
Brant (B. bernicla hrota) at Cape Vera, northern Devon 
Island (76°15'N, 89°15'W). Two days later, another sin- 
gle goose was photographed at this site, again within 
a flock of Brant, but on this occasion the bird was clear- 
ly a Cackling Goose (AJF). This bird stayed at the 
site for over a week and was attempting to establish a 
pair bond with one of the Brant. Two more Canada 
Geese were observed at Cape Vera in 2004, one single 
bird on 30 May and another on 7 June. Unfortunately, 
no data on size or flock association were recorded. 

In Quttinirpaaq National Park on northern Ellesmere 
Island, park wardens report having seen Canada Geese 
of unknown size in the vicinity of Lake Hazen (81°47'N, 
71°03'W): one lone bird on 3 June 2001 and another 
on | July 2002. 

Interestingly, Duvall and Handley (1948*) recorded 
no Canada Geese in southwestern Ellesmere Island 
in 1947, and Prach (1986*) reported none in five years 
of study at Cape Vera between 1980 and 1984. Nor 
were any Canada Geese observed during long-term 
studies on Prince Leopold Island (74°N, 90°W) from 
1988 to 2004 (Nettleship et al. 1990; A. J. Gaston, per- 
sonal communication), during aerial marine surveys 
in 1977 in Lancaster Sound (Nettleship and Gaston 
1978), during ecological research studies in the True- 
love Lowlands on northeastern Devon Island (75°41'N, 
84°35'W) in 1966-1969 and 1971-1974 (Hussell and 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Holroyd 1974; Pattie 1977), nor at Alexandra Fiord 
on eastern Ellesmere Island (78°53'N, 75°55'W) in 
annual studies since 1982 (G. Henry, personal com- 
munication). 


Discussion 

Dickson (2000b:12) noted that “As a species, Bran- 
ta canadensis is doing well and increasing rapidly in 
abundance and range’”’. The new reports and observa- 
tions presented here suggest that indeed the known 
breeding range of the Canada Goose in the eastern 
Canadian Arctic should be revised and extended at 
least 500 km further north along the northeast coast 
of Baffin Island from currently published limits. 

The breeding range of B. c. interior has long been 
known to extend to the south coast of Baffin Island 
(Palmer 1976; Bellrose 1980). We suspect that geese 
breeding along the eastern Baffin Island coast are all 
larger race, probably part of the Atlantic (B. c. interior) 
population, given that this population migrates through 
this region to Greenland (Scribner et al. 2003). Canada 
Geese reported breeding on Bylot Island (Lepage et 
al. 1998) are thought to include both Atlantic Popula- 
tion geese as well as Cackling Geese, the latter presum- 
ably of the Tallgrass Prairie Population known to breed 
in central Nunavut and on western Baffin Island around 
Foxe Basin. Geese breeding in Creswell Bay are most 
likely from the latter population. The Tallgrass Prairie 
Population is believed to be increasing and these breed- 
ing records extend their range by a distance of about 
300 km to the north and 200 km to the northeast (Dick- 
son 2000b). 

Given that Canada Geese were encountered along 
many hillsides in Frobisher Bay and south from Qik- 
iqtarjuaq during our surveys, it is likely that the dis- 
tribution of geese along the eastern coast of Baffin 
Island is sparse and fragmented at least as far north as 
Clyde River. The numbers breeding along the coast 
north of Clyde River will likely never become large, 
because the Arctic Cordillera holds only a few small 
pockets of potential feeding sites for geese, on wet 
meadows at the inner ends of fiords. The greatest scope 
for further expansion may lie in the west of Baffin Is- 
land, especially in the lakes and marshes of the Gif- 
ford River basin (70°30'N, 84°W), where Greater Snow 
Geese began to breed in the 1980s, after using it as a 
moulting area since at least the 1960s. That area, out- 
side the usual range of Inuit goose hunters, though 
visited by them in winter, has not been searched for 
geese for many years. Moreover, Canada Geese have 
been observed as far north as northern Ellesmere Island, 
and thus non-breeders may inhabit suitable habitats 
across the coastal areas of Ellesmere, Devon and north- 
ern Baffin islands. 

It is likely that some Canada Geese have frequented 
these areas for many years, at least as non-breeding 
or moulting birds, as suggested by some earlier survey 
efforts and by Inuit hunters who have reported seeing 


2005 


increases in the number of Canada Geese in the past 
30 years. Because much of this area was glaciated as 
recently as 4000 years ago, Canada Geese probably 
were not in the area until long after the last glacial peri- 
od (Dickson 2000b). Suitable breeding habitat is patchi- 
ly distributed along eastern Baffin Island, as much of 
the region is part of the Arctic Cordillera and supports 
limited, suitable vegetation for goose forage. The goose 
habitat is probably similar to Greenland, where Canada 
Geese are distributed in areas with low snow cover and 
where snow disappears first in the spring (Malecki et 
al. 2000). 

Why then have goose numbers increased in these 
regions? Although many Canada Goose populations in 
North America are steadily increasing, trends in pop- 
ulation size for the Atlantic and North Atlantic Popu- 
lations have highly fluctuated in recent years (Dick- 
son 2000b). The Atlantic Population of B. c. interior, 
which is doubtfully distinguishable based on morphol- 
ogy from the North Atlantic Population (B. c. canaden- 
sis) breeding chiefly in Labrador, decreased from an 
estimated 120 000 breeding pairs in 1988 to about 
30 000 pairs in 1995. When that decrease became ob- 
vious on its wintering range, chiefly in and around 
Maryland, the American hunting season was closed for 
several years, and the season and bag limit in southern 
Québec and Ontario were reduced. Those measures 
resulted in a rapid recovery of the breeding population 
and its expansion to around 160 000 breeding pairs in 
2002 and 2003 (CWS 2003), at which time hunting 
restrictions were relaxed. That expansion seems to 
have fuelled the growth of the breeding population in 
the Kangerlussuaq region (67°N, 50°W) of western 
Greenland which has been growing since the 1970s 
(Kristiansen et al. 1999), to now more than 2600 pairs 
(Malecki et al. 2000). Intuitively, it would appear that 
this expansion may account for the recent increase on 
Baffin Island as well. It is also possible that the moult 
migrations from increasing goose populations in areas 
in the south have led to the initiation of breeding in 
new areas in the Arctic (Mowbray et al. 2002). Still, 
our recent investigation did not allow us to determine 
whether the observed extension of breeding range 
represents a recent expansion, or rather a more com- 
plete record based on more intensive research along 
this coast in the past decade. Irrespective of the cause, 
it is clear that Canada Geese breed much further north 
on eastern Baffin Island than previously reported in 
the literature. 

Cackling Geese breed abundantly on the Great Plain 
of the Koukdjuak (66°N, 73°W), where many have 
been banded in recent years (K. M. Dickson, personal 
communication), and have been seen in many other 
parts of Baffin Island, especially on the west side, north 
at least to Bernier Bay (71°N, 88°W). The Tall Grass 
Prairie Population, to which these birds belong, defies 
enumeration at any stage of its annual life cycle. It 


MALLORY, FONTAINE, AND BOYD: CANADA AND CACKLING GEESE 


487 


migrates through central North America to winter in 
south-west Texas and north-east Mexico, passing 
though staging areas used by many other populations, 
so that it is hard to tell how its numbers may be chang- 
ing. B. hutchinsii was first found breeding in west 
Greenland in 1914, but has since done so irregularly 
and in small numbers (Salomonsen 1967). It seems to 
have been greatly outnumbered there by B. c. interior 
during the latter’s recent expansion (Boertmann 1994). 
Whether B. hutchinsii and B. c. interior can share the 
same breeding areas is not clear. 

The distribution and abundance of various water- 
birds in Nunavut are changing. Inuit hunters are pro- 
viding important observations on these changes that 
have subsequently been confirmed by scientific sur- 
veys (e.g., Robertson and Gilchrist 1998; Mallory et 
al. 2003). While much of our evidence is presently 
circumstantial, the observations and oral reports doc- 
umented here lay the baseline for directed surveys to 
confirm distributional extensions. Continued efforts 
on gathering local ecological knowledge as well as 
scientific surveys need to be continued and expanded 
if these changes are not only to be detected but ex- 
plained. 


Acknowledgements 

Field studies presented here received financial and 
logistic support from Environment Canada, the Nuna- 
vut Wildlife Management Board, and the Polar Con- 
tinental Shelf Project. Thanks to the Resource Officers 
in each of the communities for verifying information, 
D. Caswell, T. Gaston, D. Haché, G. Henry, D. Hey- 
land, D. Luukkonnen, C. Machtans, R. Malecki, and 
A. Reed for providing unpublished data, and K. Dick- 
son, A. Erskine and an anonymous referee for com- 
ments on the manuscript. 


Documents Cited (marked * in text) 

Duvall, A. J., and C. O. Handley. 1948. Second wildlife 
reconnaissance of the eastern arctic. Unpublished report, 
Canadian Wildlife Service, Yellowknife, Northwest Ter- 
ritories. 

Gilchrist, H. G., M. L. Mallory, and F. R. Merkel. 2005. 
Can traditional ecological knowledge contribute to wildlife 
management? Case studies of migratory birds. Ecology 
and Society 10: 20 [online] URL: http://www.ecologyand 
society.org/vol 1 O/iss 1/art20/ 

Heyland, J. D., and H. Boyd. 1969. An aerial reconnais- 
sance of the eastern Canadian Arctic, 20-29 July 1969, in 
search of Greater Snow Geese. Unpublished report, Cana- 
dian Wildlife Service, Iqaluit, Canada. 

Prach, R. W. Editor. 1986. Cape Vera polynya project. Un- 
published report, Canadian Wildlife Service, Iqaluit, 
Canada. 


Literature Cited 

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gists’ Union Check-list of North American Birds. Auk 121: 
985-995. 

Bellrose, F. C. 1980. Ducks, geese and swans of North Amer- 
ica, third edition. Stackpole Books, Harrisburg, Pennsy]- 
vania. 

Boertmann, D. 1994. A annotated checklist to the birds of 
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Boyd, H. 1989. Geese on Rowley Island, NWT, in 1987 and 
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Boyd, H. 1999. Tolling by breeding Brent and small Canada 
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Received 14 December 2004 
Accepted 15 September 2005 


A Survey for Federally Listed Grassland Birds at First Nations Reserves 


Scott D. STEVENS and Troy I. WELLICOME! 


Canadian Wildlife Service, Environmental Conservation Branch, Prairie and Northern Region, Room 200, 4999-98" Avenue, 
Edmonton, Alberta T6B 2X3 Canada; e-mail: troy.wellicome@ ec.gc.ca 
'Corresponding Author 


Stevens, Scott D., and Troy I. Wellicome. 2005. A survey for federally listed grassland birds at First Nations Reserves, Canadian 
Field-Naturalist 119(4): 490-496. 


We surveyed native and non-native grassland for federally listed grassland bird species at Reserves and/or Treaty Land Entitle- 
ments (TLE) belonging to five First Nations in the prairie ecozone of Canada. Ferruginous Hawk (Buteo regalis — of special 
concern), Loggerhead Shrike (Lanius ludovicianus — threatened), and Long-billed Curlew (Numenius americanus — of special 
concern) were observed at five of 335, three of 361, and five of 329 survey points within their respective geographic ranges. 
Sprague’s Pipit (Anthus spragueii-threatened) was observed at 69 of 361 survey points and accounted for 84% of sites with listed 
birds. Estimates of proportions of native and non-native grassland within 400 m of survey points indicated that Sprague’s 
Pipit preferred native grassland. Despite special effort, the Burrowing Owl (Athene culicularia-endangered) was not observed, 
although a nest from a previous year was encountered. We recorded the greatest number of species and individuals on reserves 
located in mixed and moist-mixed grasslands that contained large portions of native grassland. In many instances, field assess- 
ments revealed unsuitable habitat at sites that remote-sensing data classified as suitable. 


Key Words: Burrowing Owl, Athene cunicularia, Ferruginous Hawk, Buteo regalis, Loggerhead Shrike, Lanius ludovicianus, 


Long-billed Curlew, Numenius americanus, Sprague’s Pipit, Anthus spragueii, grassland species at risk, habitat assess- 


ment, remote sensing, Canada. 


The Canadian prairies are home to more federally 
listed birds than any other region of Canada (COSE- 
WIC 2003*). With the coming into force of The 
Species at Risk Act (SARA) in 2004, the demand for 
knowledge of locations of species at risk increased in 
Canada, particularly on federal lands. Inventories for 
listed bird species have been conducted in National 
Parks (e.g., Wynn and Jensen 1998*), National Wild- 
life Areas (e.g., Haber 1995*), and on Department of 
National Defence lands (e.g., Banasch and Barry 
1998*; Dale et al. in press). More recently, surveys 
have been undertaken on Agriculture and Agri-foods 
Canada lands (e.g., G. L. Holroyd, personal communi- 
cation). However, we are aware of no surveys for listed 
species on First Nations Reserves, even though Reserves 
are included as federal lands in the context of SARA. 

Our objective was to survey Reserves and Treaty 
Land Entitlements (TLE) for grassland bird species 
listed as endangered (Burrowing Owl, Athene cunicu- 
laria), threatened (Loggerhead Shrike, Lanius ludovi- 
cianus;, Sprague’s Pipit, Anthus spragueii) or of special 
concern (Ferruginous Hawk, Buteo regalis; Long-billed 
Curlew, Numenius americanus). Ideally, such surveys 
should be conducted during the months of May and 
June, when these birds are breeding and most conspic- 
uous (see Discussion). However, because of delays in 
securing permission for surveys, four of our surveys 
occurred between late-June until mid-July and one 
could not be conducted until mid-August. Nevertheless, 
through point counts and habitat description, we were 
able to gain a first indication of the relative importance 
of these Reserves to listed grassland birds. We were 


granted permission by five First Nations to conduct 
surveys on their lands within the prairie ecozone (Fig- 
ure 1). In Alberta, we surveyed the Blood (#148) and 
Siksika (#146) Reserves. In Saskatchewan, we sur- 
veyed Nekaneet Cree TLE, Piapot Cree First Nation 
Reserve (#75) and TLE, and Assiniboine (Carry the 
Kettle Nakota Band) Reserve (#76). Here we present 
counts of listed birds observed at each of these Reserves 
or TLE. We also summarize visual estimates of habitat 
proportions at point-count locations, and summarize 
habitat at the landscape level with remote sensing data. 
Lastly, we use habitat data, current species ranges, 
knowledge of species biology, and timing of the sur- 
veys to interpret presence/absence of listed species 
and to infer the relative importance of these lands to 
conservation of grassland bird species at risk. 


Methods 
Habitat Assessment with Remote Sensing 

Classified land-cover information was obtained for 
Alberta and Saskatchewan from the Prairie Farm Reha- 
bilitation Administration’s (PFRA) land cover (Ash- 
ton 2001*). We examined the following land-cover cat- 
egories: grassland (includes both native and non-native 
grassland), cultivated, treed, shrub-covered, and wet- 
land. We used ArcView 3.2® to summarize land-cover 
types within the boundaries of each reserve. 


Surveys 

Each survey was conducted some time between 25 
June and 14 August 2003 (Table 1) by at least two of 
five observers. All observers were trained, by Canadian 


490 


2005 


. ALBERTA 


Aspen Parkland 


STEVENS AND WELLICOME: SURVEY FOR BIRDS ON FIRST NATIONS RESERVES 


49] 


SASKATCHEWAN 
30 0 30 & Kilometers 
= —— | 


1 Blood 

2  Siksika 

3 Nekaneet 

3a Nekaneet TLE 
4 Piapot 

4a Piapot TLE 

5 Assiniboine 


Moist Mixed Grassland 


Mixed Grassland 


4 Fort Qu’Appelle 


Swift Current 


egina 


FIGURE |. First Nations Reserves and Treaty Land Entitlements (TLE) that were surveyed for federally listed grassland birds in 
this study. First Nations lands are shown in relation to provincial borders, ecoregional boundaries, and nearest cities or towns. 


Wildlife Service staff, in the visual and auditory identi- 
fication of the target species and those non-target 
species that had the most potential to be confused with 
target species. All of the Reserves and TLE contain 
areas of cultivated land and treed habitat. These habitat 
types were not surveyed because they are generally 
unsuitable for the target species (Table 2). Areas were 
selected for point-count surveys only if surrounding 
land-cover was predominantly grassland (native or 
non-native). All suitable tracts of grassland were then 
covered by survey point locations that were separated 
at 800 m intervals in each cardinal direction to form a 
uniform grid. We accessed point-count stations by 
truck or ATV (All-Terrain Vehicle). At each point, 
UTM coordinates were recorded from a hand-held 
Garmin® GPS 12 XL. An individual observer con- 
ducted a three-minute passive scan, using binoculars 
to identify and record all target species. Due to the 
relative rarity of the Burrowing Owl, we also used a 
portable CD player (Citizen® JCD4109) to broadcast 
recorded breeding calls of a male Burrowing Owl 
while scanning for an additional three minutes. 
Sprague’s Pipit were identified by their territorial songs 
during the passive phase of the scan and all other 


species by direct observation with binoculars. After 
completion of the six-minute scan, we estimated per- 
cent cover of several habitat types within a 400-m 
(50.2 ha) radius: native grassland, non-native grassland, 
cultivated land, trees, shrubs, and wetlands. This habi- 
tat assessment was performed to estimate habitat com- 
position, at the same scale as the bird counts, using 
the same variables identified in the PFRA database. 
Habitat assessment in the field also enabled us to dis- 
tinguish native from non-native grassland. 


Results 
Potential Listed Bird Species and Habitat Character- 
istics at Reserves 

The Blood Reserve, surveyed 12-14 August, is at the 
western edge of the ranges of all five listed species: 
Burrowing Owl (Wellicome and Holroyd 2001), Log- 
gerhead Shrike (Yosef 1996), Sprague’s Pipit (Robbins 
and Dale 1999), Ferruginous Hawk (Bechard and 
Schmutz 1995), and Long-billed Curlew (Dugger and 
Dugger 2002). The Blood was the largest reserve in 
this study (Table 2), with 40% of overall land-cover 
classified as grassland and most of the remaining land 
classified as cultivated (Table 2). Remote sensing 


492 THE CANADIAN FIELD-NATURALIST Vol. 119 


TABLE |. Summary of listed grassland birds at First Nations Reserves and Treaty Land Entitlements (TLE) during 2003 point- 
count surveys. BUOW = Burrowing Owl; FEHA = Ferruginous Hawk; LBCU = Long-billed Curlew; LOSH = Loggerhead 
Shrike; SPPI = Sprague’s Pipit. Numbers for each species denote survey points at which they were detected. In parentheses 
is the total number of individuals detected. 


Reserve Ecoregion Potential # Date of # Survey BUOW FEHA LBCU LOSH SPPI 
listed species Survey Points 

Blood Moist-mixed Grassland 5 12-14 August 96 0 | | 0 
Siksika Moist-mixed Grassland 5 25-27 June 148 O* D, 4 (6) 3 44(2) 
Nekaneet TLE Mixed Grassland 5) 9-11 July 85 0 2 0 0 227 
Piapot TLE Moist-mixed Grassland 4 7-8 July 6 0 0 — 0 2 
Piapot Aspen Parkland ye 7-8 July 1) — — — 0 1 
Assiniboine Aspen Parkland 2 3-4 July 1] — — — 0 0 
Total 361 0 5) o(7) 3. ‘69:(83) 


*One inactive Burrowing Owl nest site, which appeared to have been occupied in previous years, was noted at the Siksika 
reserve. 


TABLE 2. Total hectares and %cover of habitat types at First Nations Reserves and Treaty Land Entitlements (TLE) based on 
remote sensing. In parentheses, total number of hectares surveyed at each reserve and summary of habitat percentages with- 
in a 400-m radius of survey-points. Percentages of habitat types at survey-points reported as mean + standard error. For 
“Grassland”, %Native and %Non-native was distinguished visually in the field. 


Reserve Total in ha %Grassland %Cultivated % Trees %Shrubs% %Wetlands 
(%Native) (%Non-native) 
Blood 140124 40.3 58.9 0.2 0.1 <0. 1 
(4823 ) (77.8 = 2.6) (O83 =.0:4).. -Gi933 = 25) (0.302) (420.1) (L503) 
Siksika 71339 68.1 en 0.9 5 0.0 
(7436 ) (88.8 + 1.6) (2.3 + 0.9) (37 = 8.3) (0.9+ 0.4) (0.7 20.2) (1.3202) 
Nekaneet TLE 10725 95.2 2.0 0.1 ey 0.0 
(4271 ) (Gore 32) “Iso 23) Cw i GA+ 0.9) 63207) (Et=02) 
Piapot TLE 11244 2.6 91.0 0.3 1.9 32 
(301 ) (23.G'25, 5)" (CALWECSS) * (45921050) 6324.9)" 905402) (0.7 22038) 
Piapot 8140 38.4 44.4 7.4 2.2 5:2 
(754 ) (0.3 =0.3) (67.7 +10.4) (8.3 + 8.3) (23.0'+10.6) (5240.2) (0.2402) 
Assiniboine 16375 13 19.4 3.9 0.0 0.3 
(553') (14.0 +6.2) (65.0 +10.8) (O -=10) (iS.7 = 5:0) Cs) (277 = 14) 
Overall 259197 54.6215 405 2129 54 = 34 1.4+ 0.4 1.6+0.8 
(18138 ) (74.7 = 16) Ot 1.2) (O42 14) (2.22 0.5) (:0+033) (3520) 


attributed very little land cover to trees and shrubs. 
Overall land-cover estimates for trees and shrubs were 
similar to what we observed at survey-points (Table 2). 
Visual estimates at point-count locations revealed that 
most grassland is native. 

The Siksika Reserve is also near the western edge 
of the ranges of all target species. We surveyed the 
Siksika Reserve 25-27 June (Table 1). It was the sec- 
ond largest of the five reserves (Table 2), with 68% of 
overall land-cover classified as grassland and scarce 
tree and shrub cover (Table 2). Visual estimates at 
point-count locations revealed that grasslands were 
almost exclusively native. Tree and shrub cover at 
survey-points was similar to overall land-cover esti- 
mates (Table 2). 

Located northeast of Cypress Hills, the Nekaneet 
Cree Nation Reserve contains rolling habitat that is 
extensively treed and thus unsuitable for grassland 
species. However, Nekaneet TLE, which is separate 


from the reserve proper (Figure 1), 1s within the ranges 
of all of the target species and has suitable habitat. 
On Nekaneet TLE, remote sensing classified 95% of 
land-cover as grassland, with very minor shrub and 
cultivation coverage (Table 2). However, visual esti- 
mates from survey points revealed that as much as 
15% of the grassland is non-native and that trees and 
shrubs are in somewhat higher proportions within the 
grasslands than is indicated by remote sensing. 
Piapot Reserve is within the range of the Logger- 
head Shrike and Sprague’s Pipit and is near the east- 
ern edge of the Burrowing Owl and Ferruginous Hawk 
ranges, at the transition from Moist Mixed Grassland 
to Aspen Parkland. The Reserve was surveyed 7-8 
July (Table 1). Although remote sensing classified 
38% of the Reserve as grassland, visual estimates 
indicated that very little native grassland remains on 
the Reserve, most having been converted to non- 
native pasture. These pastures contained a much higher 


2005 


STEVENS AND WELLICOME: SURVEY FOR BIRDS ON FIRST NATIONS RESERVES 493 


TABLE 3. Summary of habitat characteristics for points at which listed species were observed. Percentages reported as mean + 
standard error. The category ‘Present’ includes locations where any of the five listed species was observed. “Absent” includes 


all locations at which listed species were not observed. 


Species N %Native %Tame %Cultivated %Tree %Shrub % Wetlands 
FEHA 5 ONG. z 3.2 0+0 2.0 + 2.0 0.4 + 0.4 D2 2.0 4.8+3.9 
LBCU 2) 17.4 = 13.3 0+0 21.6 + 13.4 0+0 0+0 LO 03 
LOSH 3 7a OS 0+0 0+0 1.0+0.6 [30.7 O32 0:3 
SPPI 69 S71 £23 1.8210 3.5: 1.5 Stl 4.0+ 1.4 L202 
Present 82 86.2 + 2.4 Loz 09 4.6+ 1.6 iMac =s Uae, 3.7410 1.4+0.3 
Absent 279 MALO ALD Pee 1220 3 2.4 + 0.6 L735 20:3 15 10-2 


FEHA -— Ferruginous Hawk; LBCU — Long-tailed Curlew; LOSH — Loggerhead Shrike; SPPI — Sprague’s Pipit. 


proportion of trees than is indicated by remote sens- 
ing (Table 1). 

Piapot TLE, south and west of the Piapot Reserve 
(Figure 1), is well within Moist Mixed Grassland but 
east of the range of the Long-billed Curlew. Ninety-one 
percent of the land is cultivated, and visual estimates 
indicated that remaining grassland (3%) contains more 
non-native than native grassland. Trees and shrubs 
were in higher proportion within the areas surveyed 
than is indicated by remote sensing (Table 2). 

Assiniboine Reserve is located in the Aspen Park- 
land, outside of the current ranges of the Burrowing 
Owl, Ferruginous Hawk, and Long-billed Curlew, but 
inside the range of both the Loggerhead Shrike and 
Sprague’s Pipit. Remote sensing classified 76% of 
land cover in the Assiniboine Reserve as grassland 
(Table 2). Visual estimates at survey-points, however, 
revealed that 65% of grassland was non-native and that 
trees and shrubs were in higher proportions than indi- 
cated by remote sensing. 


Species Observations 

Listed birds were observed at 82 of the 361 (23%) 
survey points (Table 1). Of the 82 points with listed 
birds, Sprague’s Pipit was observed at 69 (84%); it 
was relatively common at Siksika (44 of 148 points; 
30%), Nekaneet TLE (21 of 85 points; 25%), and 
Piapot TLE (two of 6 points; 33%), but was seldom 
detected at the Blood (1 of 96 points; 1%) and Piapot 
(one of 15 points; 7%) Reserves. Ferruginous Hawks 
were observed at one location on the Blood Reserve 
and two locations on the Siksika Reserve and Nekaneet 
TLE, but were not detected at Piapot TLE, though this 
land is within their geographic range. Long-billed 
Curlews were most abundant at Siksika Reserve (four 
of 148 points; six individuals; Table 1) but were 
encountered only at one other location, on the Blood 
Reserve. Loggerhead Shrikes were observed only at 
Siksika (three of 148 points; 2%). 

Despite special effort to locate them (i.e., taped 
calls), the Burrowing Owl was the only listed species 
not observed on any of the Reserves. However, we did 
find two burrows (10 m apart) at Siksika that we 
judged, based on several small rodent skeletal remains 


on and near the mounds, had previously been a Bur- 
rowing Owl nest and roost. There were no ow! pellets 
or whitewash, and burrow entrances were beginning to 
collapse, indicating the site had not been used for at 
least one year (see California Burrowing Owl Consor- 
tium 1997). 

To infer habitat selection for Sprague’s Pipits, we 
compared visual estimates of percentage of native 
grassland at point locations where pipits were present 
versus where they were absent. We considered only 
Nekaneet TLE and Siksika, where the species was 
observed in highest abundance and where timing of 
surveys was concurrent with territorial songs (Rob- 
bins 1998). At both Nekaneet TLE and Siksika, the 
percentage of native prairie was higher at locations 
where Sprague’s Pipits were present (Table 3). Al- 
though sample sizes were small, Ferruginous Hawks, 
Long-billed Curlews, and Loggerhead Shrikes showed 
a similar pattern (Table 3). 


Discussion 

The timing of these surveys most likely reduced 
detection rates of some target species, particularly the 
August survey at the Blood Reserve. Although Bur- 
rowing Owls typically begin leaving Canada on south- 
ward migration during September and October (Todd 
et al. 2003), males are most aggressive and conspicu- 
ous during egg laying and brood rearing (May through 
June) and thus more likely to respond to call play- 
backs during that time (Haug and Didiuk 1993; Shyry 
et al. 2001). Long-billed Curlews depart from northern 
parts of their breeding range by mid-July and from all 
parts of their breeding range by mid-August (Dugger 
and Dugger 2002). Sprague’s Pipits perform persistent 
displays and songs from mid-April through the third 
week of May, with another period of elevated display 
from mid-June to mid-July; however, their singing 
ceases almost entirely by mid-August (Robbins 1998). 
Ferruginous Hawks (Bechard and Schmutz 1995) and 
Loggerhead Shrikes (Yosef 1996) do not begin migra- 
tion until September, and these species may be most 
easily detected during breeding in May and June. 
Though timing for the surveys in this study was sub- 
optimal, the relative magnitude of our counts of listed 


494 


grassland birds corresponded roughly to overall esti- 
mates of abundance for each species in Canada. 

The Burrowing Owl is listed as endangered in Can- 
ada (Wellicome and Haug 1995*; COSEWIC 2003*) 
and has been essentially extirpated from Manitoba. 
The range of this species within southern Alberta and 
Saskatchewan is currently half the size it was 20-30 
years ago (Wellicome and Holroyd 2001). Fewer than 
800 pairs are estimated to remain on the Canadian 
Prairies (Todd 2005*), and our survey provides support 
for the rarity of this species. Like many areas across 
the Canadian prairies, the Blood and Siksika reserves, 
and Nekaneet TLE, appear to contain suitable but 
vacant Burrowing Owl habitat. 

The Ferruginous Hawk, listed as a species of special 
concern in Canada (COSEWIC 2003*), still occupies 
many areas where it was reported in historical times, 
but has declined at the northern edge of its range in 
Alberta, Saskatchewan, and Manitoba (Schmutz 1995*; 
Schmutz et al. 1994*). The breeding population in 
Canada is estimated at 2000-4000 pairs (Schmutz 
1995*; Schmutz et al. 1994*). Although Ferruginous 
Hawks prefer elevated nest sites, such as isolated trees 
or platforms, they occur in large tracts of open, gener- 
ally arid habitats dominated by grasses or sagebrush 
(Bechard and Schmutz 1995). Schmutz (1989) showed 
that they were more abundant where levels of cultiva- 
tion were low. Our results agree with these previous 
studies, as Ferruginous Hawks were found on reserves 
with the largest areas of native grassland. 

Prairie populations of the Loggerhead Shrike, listed 
in Canada as threatened (Cadman 1985*; COSEWIC 
2003*), have declined in numbers since the 1960s 
(Yosef 1996), and the northern limit of their range has 
retracted (Telfer et al. 1989). The most recent popula- 
tion estimate is 2500 pairs in Alberta, 7000 pairs in 
Saskatchewan, and 500 pairs in Manitoba (Johns et al. 
1994*). All of the Reserves we surveyed were within 
the geographic range of Loggerhead Shrikes, but we 
found shrikes only at Siksika. Siksika contains spo- 
radic shrub and tree cover, which Loggerhead Shrikes 
require for nesting habitat (Yosef 1996), and large 
areas of grassland to support prey species of Logger- 
head shrikes. Although their abundance is correlated 
with availability of pastureland (Gawlik and Bildstein 
1993), Loggerhead Shrikes occur at relatively low 
densities (Telfer 1993) and thus may be more diffi- 
cult to detect than Long-billed Curlews, Ferruginous 
Hawks, and vocalizing Sprague’s Pipits. 

Long-billed Curlews, listed as a species of special 
concern in Canada (De Smet 1992*), are now consid- 
ered extirpated in Manitoba (De Smet 1992*; Hill 
1998*) and very rare in areas of southeastern Sask- 
atchewan, where they were once common (Renaud 
1980). Although the population in Alberta has been 
estimated at approximately 24 000 (assuming equal sex 
ratio; Saunders 2001*), it is suspected they are declin- 
ing rapidly in eastern parts of their range, including 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Saskatchewan (De Smet 1992*; Saunders 2001*). 
Long-billed Curlews nest close to wetter areas in short 
grass or mixed-grassland habitat (Hooper and Pitt 
1996*) and avoid areas with trees and high densities of 
shrubs (Pampush and Anthony 1993). We did not 
observe Long-billed Curlews at Nekaneet TLE, which 
is within their geographic range, perhaps because the 
grasslands there contain few wetlands and relatively 
high shrub-cover. Habitat characteristics were general- 
ly similar between Blood and Siksika Reserves, but we 
observed more Long-billed Curlews at Siksika. One 
difference that may help explain this result is the lower 
proportion of wetlands at the Blood Reserve. Perhaps 
more importantly, however, our survey at the Blood 
Reserve was late, so many Long-billed Curlews may 
have already started to migrate south. 

Sprague’s Pipit is listed as threatened in Canada 
(COSEWIC 2003*). Although still locally abundant, 
populations of Sprague’s Pipit have been declining rap- 
idly in parts of their range, with the greatest decline 
occurring in Canada’s prairie provinces (Sauer et al. 
1997*). Populations in Alberta and Saskatchewan, 
where highest densities occur, have declined by 9.4% 
and 5.4% per year, respectively (Prescott and Davis 
1998*). Sprague’s Pipits prefer native grassland, rarely 
being found in cultivated fields or where native grasses 
have been replaced by introduced species (Robbins and 
Dale 1999). Accordingly, the only Reserve where 
Sprague’s Pipits were not observed in this study was 
the Assiniboine, which is composed predominantly of 
non-native grassland. The species was found in relative 
abundance at the Nekaneet TLE and the Siksika 
Reserve, which both contain substantial proportions of 
native grassland. The Blood Reserve, which also con- 
tains large tracts of native grassland, albeit at the west- 
ern edge of the range, probably contains a larger popu- 
lation of Sprague’s Pipit than our late survey would 
suggest. By mid-August male territorial vocalizations 
are rare (Robbins 1998) and thus Sprague’s Pipit are 
exceedingly difficult to detect at that time. 

Our comparison of percentage native grassland be- 
tween those sites at Siksika and Nekaneet TLE where 
Sprague’s Pipit were and were not detected (in surveys 
concurrent with male territorial songs) support other 
studies showing a preference for native grassland by 
this species (Robbins and Dale 1999). Although sample 
sizes were small, Ferruginous Hawks, Long-billed 
Curlews, and Loggerhead Shrikes were also observed 
at point locations with high percentages of native 
prairie, suggesting the importance of native grasslands 
to these species. 

The PFRA land-cover data can be misleading for 
large-scale habitat assessments because they do not dis- 
tinguish native from non-native grassland, and tend to 
underestimate shrub and tree cover within grasslands. 
Based on visual habitat estimates, the Assiniboine and 
Piapot Reserves, which are both in aspen parkland, 
contained high proportions of trees and tame pasture 


2005 


and were of relatively little importance to listed grass- 
land birds. In contrast, the Siksika and Blood Reserves, 
which are in moist-mixed grassland, and Nekaneet 
TLE, which is in mixed grassland, each contain rela- 
tively large proportions of native grassland and appear 
to be important areas for listed grassland birds. 


Acknowledgments 

Funding support for this project was provided by the 
Canadian Wildlife Service, in the Prairie and North- 
ern Region of Environment Canada. We extend our 
thanks to Dave Duncan, who encouraged us to under- 
take these surveys, and to the Blood, Siksika, Nekaneet 
Cree, Piapot Cree, and Carry the Kettle First Nations 
for granting us access to their lands. For excellent assis- 
tance in the field, we thank Stephanie Grossman, David 
Junor, and Joann Skilnick. We also thank Geoffrey L. 
Holroyd, of the Canadian Wildlife Service, for sharing 
a personal communication about his species-at-risk 
surveys on Agriculture and Agri-foods Canada lands. 
Special thanks to David Junor for using the PFRA 
GIS dataset to calculate proportions of land-cover 
within each reserve and also for preparing Figure 1. 


Documents Cited [marked * in text] 

Ashton, J. 2001. PFRA’s Generalized Land Cover, Version 1. 
Prairie Farm Rehabilitation Administration, Agriculture 
and Agri-foods Canada, Regina, Saskatchewan. http://www. 
agr.gc.ca/pfra/gis/. 

Banasch, U., and S. J. Barry. 1998. Raptor component 
report: Canadian Forces Base Suffield National Wildlife 
Area wildlife inventory. Unpublished Canadian Wildlife 
Service Report, Edmonton, Alberta. 51 pages. 

Cadman, M. D. 1985. Status report on the Loggerhead Shrike 
in Canada. Committee on the Status of Endangered Wild- 
life in Canada, Ottawa, Ontario. 96 pages. 

COSEWIC. 2003. Canadian Species at Risk, May 2003. 
Committee on the Status of Endangered Wildlife in Can- 
ada, Ottawa, Ontario. 43 pages. 

De Smet, K. D. 1992. Status report on the Long-billed Curlew 
Numenius americanus in Canada. Committee on the Status 
of Endangered Wildlife in Canada, Ottawa, Ontario. 44 
pages. 

Haber, E. 1995. Species at Risk and invasive plants of 
National Wildlife Areas and Migratory Bird Sanctuaries. 
Unpublished Canadian Wildlife Service Report, Ottawa, 
Ontario. 170 pages. 

Hill, D. P. 1998. Status of the Long-billed Curlew (Nume- 
nius americanus) in Alberta. Alberta Environmental Pro- 
tection, Fisheries and Wildlife Management Division, and 
Alberta Conservation Association, Wildlife Status Report 
(16), Edmonton, Alberta. 20 pages. 

Hooper, T. D., and M. D. Pitt. 1996. Breeding bird commu- 
nities and habitat associations in the grasslands of the Chil- 
cotin Region, British Columbia. Canada-British Columbia 
Partnership Agreement on Forest Resource Development: 
FRDA IL. 

Johns, B., E. Telfer, M. Cadman, D. Bird, R. Bjorge, K. 
De Smet, W. Harris, D. Hjertaas, P. Laporte, and R. 
Pittaway. 1994. National Recovery Plan for the Logger- 
head Shrike. Recovery of Nationally Endangered Wild- 
life Report number 7, Ottawa, Ontario. 32 pages. 


STEVENS AND WELLICOME: SURVEY FOR BIRDS ON FIRST NATIONS RESERVES 


495 


Prescott, D. R. C., and S. K. Davis. 1998. Status Report on 
the Sprague’s Pipit Anthus spragueii in Canada. Status 
Report to the Committee on the Status of Endangered 
Wildlife in Canada, Ottawa, Ontario. 33 pages. 

Sauer, J. R., J. E. Hines, G. Gough, I. Thomas, and B. G. 
Peterjohn. 1997. The North American breeding bird sur- 
vey results and analysis, Version 96.4. Patuxent Wildlife 
Research Center, Laurel, Maryland. http://www.mbr-pwrc. 
usgs.gov/bbs/bbstext.html 

Saunders, E. J. 2001. Population estimate and habitat asso- 
ciations of the Long-billed Curlew (Numenius americanus) 
in Alberta. Alberta Fish and Wildlife Division, Species at 
Risk Report number 25, Edmonton, Alberta. 57 pages. 

Schmutz, J. K. 1995. Updated Status Report on the Ferrug- 
inous Hawk, Buteo regalis, in Canada. Committee on the 
Status of Endangered Wildlife in Canada, Ottawa, Ontario. 
18 pages. 

Schmutz, J. K., S. H. Brechtel, K. D. De Smet, D. G. 
Hjertaas, C. S. Houston, and G. L. Holroyd. 1994. 
National Recovery Plan for the Ferruginous Hawk. Report 
number 11. Recovery of Nationally Endangered Wildlife 
Committee, Ottawa, Ontario. 35 pages. 

Todd, L. D. 2005. Status of the Burrowing Owl (Athene cuni- 
cularia) in Alberta: update 2005. Alberta Sustainable Re- 
source Development, Wildlife Status Report Number 11 
(Update 2005), Edmonton, Alberta. 28 pages. 

Wellicome, T. I., and E. A. Haug. 1995. Second update of 
status report on the Burrowing Owl Speotyto cunicularia 
in Canada. Committee on the Status of Endangered Wild- 
life in Canada, Ottawa, Ontario. 23 pages. 

Wynn, M., and O. Jensen. 1998. Sage grouse monitoring, 
1997. Grasslands National Park Annual Report Volume 3: 
54-56. 


Literature Cited 

Bechard, M. J., and J. K. Schmutz. 1995. Ferruginous Hawk 
(Buteo regalis). In The Birds of North America (172). Edit- 
ed by A. Poole and F. Gill. The Academy of Natural Sci- 
ences, Philadelphia, Pennsylvania; The American Ornith- 
ologists’ Union, Washington, D.C. 

California Burrowing Owl Consortium. 1997. Burrowing 
owl survey protocol and mitigation guidelines. Raptor Re- 
search Reports 9: 171-177. 

Dale, B. C., M. R. Norton, P. S. Taylor, and J. P. Goossen. 
In press. Protecting the regional avifauna-topographic, 
habitat, and bird diversity in a prairie reserve. Canadian 
Field-Naturalist. 

Dugger, B. D., and K. M. Dugger. 2002. Long-billed Curlew 
(Numenius americanus). In The Birds of North America, 
(62B). Edited by A. Poole and F. Gill. The Academy of 
Natural Sciences, Philadelphia, Pennsylvania; The Amer- 
ican Ornithologists Union, Washington, D.C. 

Gawilik, D. E., and K. L. Bildstein. 1993. Seasonal habitat 
use and abundance of loggerhead shrikes in South Caroli- 
na. Journal of Wildlife Management 57: 352-357. 

Haug, E. A., and A. B. Didiuk. 1993. Use of recorded calls 
to detect burrowing owls. Journal of Field Ornithology 64: 
188-194. 

Pampush, G. J., and R. G. Anthony. 1993. Nest success, 
habitat utilization, and nest-site selection of Long-billed 
Curlews in the Columbia Basin, Oregon. Condor 95: 957- 
967. 

Renaud, W. E. 1980. The Long-billed Curlew in Saskatche- 
wan: status and distribution. Blue Jay 38: 221-237. 


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Robbins, M. R. 1998. Display behavior of male Sprague’s 
Pipits. Wilson Bulletin 110: 435-438. 

Robbins, M. B., and B. C. Dale. 1999. Sprague’s Pipit 
(Anthus spragueii). In The Birds of North America (439). 
Edited by A. Poole and F. Gill. The Academy of Natural 
Sciences, Philadelphia, Pennsylvania; The American Or- 
nithologists’ Union, Washington, D.C. 

Schmutz, J. K. 1989. Hawk occupancy of disturbed grass- 
lands in relation to models of habitat selection. Condor 
91: 362-371. 

Shyry, D. T., T. I. Wellicome, J. K. Schmutz, G. L. Erick- 
son, D. L. Scobie, and R. F. Russell. 2001. Burrowing 
Owl population-trend surveys in southern Alberta: 1991- 
2000. Journal of Raptor Research 35: 310-315. 

Telfer, E. S., C. Adam, K. De Smet, and R. Wershler. 1989. 
Status and distribution of the Loggerhead Shrike in western 
Canada. Canadian Wildlife Services Progress Notes Num- 
ber 184. 4 pages. 


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Telfer, E. S. 1993. Habitat change as a factor in the decline 
of the western Canadian Loggerhead Shrike, Lanius ludo- 
vicianus, population. Canadian Field-Naturalist 106: 321- 
326. 

Todd, L. D., R. G. Poulin, T. I. Wellicome, and R. M. 
Brigham. 2003. Post-fledging survival of Burrowing Owls 
in Saskatchewan. Journal of Wildlife Management 67: 
512-519. 

Wellicome, T. I., and G. L. Holroyd. 2001. The second 
international Burrowing Owl Symposium: background 
and context. Journal of Raptor Research 35: 269-273. 

Yosef, R. 1996. Loggerhead Shrike (Lanius ludovicianus). 
In The Birds of North America (231). Edited by A. Poole 
and F. Gill. The Academy of Natural Sciences, Philadel- 
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Washington, D.C. 


Received 2 February 2004 
Accepted 27 October 2005 


Addition to the Flora of Canada? A Specimen from the Arctic 
Archipelago, Northwest Territories Links Two Allopatric Species 
of Alkali Grass, Puccinellia 


L. L. CONSAUL, L. J. GILLESPIE, and K. I. MACINNEs! 


Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada; email: lconsaul @ mus-nature.ca; 
lgillespie @ mus-nature.ca. 
' Box 21, Site B, 1052 Cedarwood Drive, Verona, Ontario KOH 2WO Canada; email: macinnes @kingston.net 


Consaul, L. L., L. J. Gillespie, and K. I. MacInnes. 2005. Addition to the flora of Canada? A specimen from the Arctic Archipel- 
ago, Northwest Territories links two allopatric species of alkali grass, Puccinellia. Canadian Field-Naturalist 119(4): 
497-506. 


A single herbarium specimen from Banks Island in the Canadian National Herbarium, Ottawa, is closest to Puccinellia wrightii 
(Puccinellia sect. Pseudocolpodium). This would represent a species new to Canada and an extension of over 1100 km from 
the previously known range in NW Alaska and NE Russia. The morphological characteristics of this specimen are com- 
pared with all taxa in P. section Pseudocolpodium and the North American P. arctica aggregate. Principal components analysis 
supports placement of this specimen in P. section Pseudocolpodium near P. wrightii, where it contributes to a morphological 
continuum between this species and P. vahliana. The new combination Puccinellia wrightii var. flava is made and a map of 
the current known distribution of the species in P. section Pseudocolpodium is presented. 


Key Words: Alkali grass, Puccinellia wrightii, Puccinellia section Pseudocolpodium, principal components analysis, Canadian 


Arctic Archipelago. 


Alkali grasses (Puccinellia Parl.) are tufted, pioneer- 
ing grasses which are usually found along the seashore 
or on interior salt pans and alkaline marine sediments 
in both temperate and arctic regions. They have soft, 
often inrolled leaves, panicles with several florets per 
spikelet, small glumes that are shorter than the awnless 
lemmas, and lemmas with rounded backs. Puccinellia 
is currently the largest Arctic grass genus. 

Puccinellia has a long history of taxonomic uncer- 
tainty and plants are often difficult to identify. Fernald 
and Weatherby (1916, page 1) called it “even to agros- 
tologists, one of the most perplexing groups of grass- 
es...,’ Polunin (1940, page 67) commented that: “the 
characters are variable... [P. angustata] is a most 
unsavoury aggregate...,” and Davis (1993, page 202) 
called it “one of the more controversial genera in the 
grass family in terms of species delimitation.” Where- 
as over 300 species have been described worldwide, 
30 to 80 species have been accepted over the span of 
the last 20 years (Gould and Shaw 1983; Davis 1983; 
Watson and Dallwitz 1999°). 

During examination of specimens for morphologi- 
cal studies on alkali grasses of the Canadian Arctic 
Archipelago (CAA) (Consaul and Gillespie 2001) and 
the grasses of the CAA (Aiken et al. 2000*), a herbari- 
um specimen was found that did not key to any known 
Canadian grass species and was not discussed in either 
of these works. It had been collected by Kaye 
MaclInnes during an ecological survey with John Lam- 
bert (Carleton University) of an area on Banks Island 
near Egg River. This area was under study because it 


* See Documents Cited section 


was proposed as an International Biological Program 
(IBP) site. Although not designated as an IBP, the area 
falls within the Banks Island No | Migratory Bird 
Sanctuary, which protects the nesting habitat of the 
Lesser Snow Goose, Chen caerulescens caerulescens. 
Knowledge of baseline botanical information in this 
area, especially from specimens in the 1970s, is of 
interest because Snow Goose colonies have increased 
dramatically in recent years (McRae et al. 1994). At 
the Banks Island sanctuary the geese are now foraging 
further inland and upland than their original grounds 
in the valley of the Big River (Jim Hines, Natural 
Resources Canada, personal communication, 2003). 
While geese feed commonly on P. phryganodes, a 
stoloniferous species that grows in low saline environ- 
ments in the area, the Puccinellia specimen in question 
was growing on upland slopes where the geese have 
been spreading. 

Puccinellia in the North American Arctic may be 
placed into three sections of the classification by 
Tzvelev (1976). Puccinellia section Paralochloa con- 
tains the stoloniferous species P. phryganodes (Trin.) 
Scribn. & Merr. Puccinellia section Puccinellia, con- 
taining most species in the CAA, has glumes that are 
relatively uneven in size, with the first glume being 
less than 2 the length of the first lemma, and usually 
short anthers. The P. arctica aggregate with long 
anthers, first recognized by Polunin (1959) and 
endemic to North America, likely belongs in this sec- 
tion but has not been formally placed here because the 
sectional classification was established for species in 


497 


498 


Russia. Puccinellia section Pseudocolpodium has 
glumes that are relatively even in size, with the first 
glume usually over 2 the length of the first lemma, 
fairly long anthers, and thick, crinkled roots. World- 
wide, six species have been described in this section: 
P. beringensis Tzvelev, P. byrrangensis Tzvelevy, P. col- 
podioides Tzvelev, P. jenisseiensis (Roshev.) Tzvelev, 
P. vahliana (Liebm.) Scribn. and Merr., and P. wrightii 
(Scribn. & Merr.) Tzvelev. The last two species, of 
which the latter is represented by two varieties (see 
under Taxonomy below), were originally described in 
the genus Colpodium. Puccinellia vahliana is the only 
Canadian species; P. wrightii is from Alaska and Rus- 
sia and is the next closest geographically; and the 
other species are from Russia. 

Previous determinations by S. G. Aiken (Canadian 
Museum of Nature) had placed the Banks Island speci- 
men in Puccinellia arctica (Hooker) Fernald and 
Weath. (in 1989), and later in P. agrostidea T. J. 
S¢rensen (in 1994) of the P. arctica aggregate owing 
to its long anthers and fairly uneven glumes. In early 
principal components analyses for the morphological 
study of Puccinellia by Consaul and Gillespie (2001), 
this specimen did not associate with any groups. Upon 
reexamination of the specimen, LLC identified it (in 
2000) as “P. vahliana x ?” of section Pseudocolpodi- 
um because it had relatively long glumes and thick, 
crinkled roots. 

This paper presents the findings of our investigation 
into the identity of this specimen. The specific objec- 
tives are (1) to confirm whether this taxon belongs to 
the large-anthered P. arctica aggregate, or to P. sec- 
tion Pseudocolpodium, and (2) to determine whether it 
aligns with a currently described species in the genus. 


Methods 

A single specimen from the National Herbarium of 
Canada (N.W.T., Banks Island, Egg River, Area #16, 
K. L. MacInnes s.n., 24 July 1971, CAN 535850) is 
the subject of this study [collection acronyms follow 
Holmgren et al. 1990]. We consulted floras of Canada, 
Alaska, and Russia to determine the putative identifi- 
cation of the specimen. After preliminary screening 
the specimen was considered to be most closely relat- 
ed to species in Puccinellia section Pseudocolpodium 
or to members of the North American P. arctica aggre- 
gate (P. agrostidea, P. arctica, and P. poacea). We com- 
pared the morphological characteristics of the CAN 
535580 specimen with specimens of these species. 

For P. agrostidea (including holotype), P. arctica, P. 
poacea (including holotype), and P. vahliana, many 
specimens were initially examined to determine the 
range of variation, and a set of ten to 12 specimens of 
each (Appendix 1) were scored (Table 1). Puccinellia 
agrostidea was excluded from Table 1 because of the 
uncertainty of its taxonomic limits, as well as its rela- 
tively small florets and anthers. Eight specimens of P. 
wrighti var. wrightii, including the holotype, were 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


obtained and examined. Material was scant or lacking 
for the other taxa. Only three specimens of P. colpodi- 
oides were available, and the holotype of Pwrightii 
var. flava (the only collection known to date) was 
included. Measurements were made on five separate 
culms of both the CAN 535850 and the P. wrightii var. 
flava (Scribn. & Merr.) Consaul (see Taxonomy sec- 
tion below) collections, and each culm was considered 
as a separate Operational Taxonomic Unit (OTU), 
because we had only a single herbarium sheet of each. 
Data from original descriptions of P. beringensis, P. 
byrrangensis, and P. jenesseiensis were used for com- 
parison since no specimens were available. Of these, 
only P. byrrangensis was included in the final dataset, 
since initial examination showed that CAN 535850 
keyed out close to this species but not to the other two. 

The characters examined were the same as those 
found useful in separating Puccinellia taxa in the 
analyses of Consaul and Gillespie (2001: Tables 2 and 
4). Two additional qualitative characters, “presence or 
absence of hyaline margin” and “inflorescence open or 
contracted,” were added since they had been used in 
keys to these species. For P. wrightii and P. colpodi- 
oides, we kept the data of the non-type herbarium 
specimens separate from the data of the protologue 
and the holotype. For P. byrrangensis only data from 
descriptions was used. 

A principal components analysis (PCA) was per- 
formed using SYSTAT 7.0 on all taxa for which we 
had specimens, except one. Puccinellia wrightii var. 
flava was excluded because its very large spikelet 
measurements make it clearly different from CAN 
535850. A subset of five representative specimens of 
most taxa, the three of P. colpodioides, and the five 
culms of CAN 535850 were used for the PCA to bal- 
ance the ordination because of the low number of 
specimens of some species available for analysis. We 
recognized that the dispersion of data points in any 
ordination analyses we performed on these measure- 
ments for the single collection of CAN 535850 (likely 
representing a single plant) might be narrower than the 
dispersion when each OTU was from a separate pop- 
ulation. 

The PCA was performed on the quantitative char- 
acters listed in Table 1, plus height, ligule length, 
length of first internode of panicle, palea length, callus 
hair length, extent of hair from base of lemma, and 
extent of hair from apex of palea (characters from 
Consaul and Gillespie 2001). A dataset reduction pro- 
cedure used by Davis (1983) and established as use- 
ful in Consaul and Gillespie (2001) was used here. In 
this procedure, a group of OTUs which separated 
from the rest of the data was removed before the 
remaining reduced dataset was analyzed in a subse- 
quent PCA. In this way, the PCA would calculate the 
axes on the variation present in the remaining smaller 
group, thus potentially revealing further groupings 
among the smaller set of OTUs. 


499 


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CONSAUL, GILLESPIE, AND MACINNES: ALKALI GRASS FROM THE ARCTIC ARCHIPELAGO 


50] 


FicureE |. Map of species in Puccinellia section Pseudocolpodium. Star in circle = CAN 535850. Solid circles and dot pat- 
terns = P. vahliana; open squares = P. wrightii; solid square = P. beringensis, representing several localities; open 
triangle = P. colpodioides; diagonal hatching = P. byrrangensis; horizontal hatching = P. jenisseiensis; solid triangle 


= P. wrightii var. flava. 


The current distribution of the species in P. section 
Pseudocolpodium was mapped (see Figure 1). The 
distribution of P. vahliana is based on specimens from 
CAN and DAO (solid circles), as well as the maps in 
Hultén (1968) (area indicated by dot pattern) and 
Cody (1996) (Yukon record). The open squares repre- 
senting P. wrightii var. wrightii are based on specimens 
from CAN, DAO, ALA and US, as well as the dot 
map in Probatova (1985) and text in Tzvelev (1964). 
The open triangle of P. colpodioides is based on several 
specimens from CAN and ALA, and text in Tzvelev 
(1964). The solid triangle representing P. wrightii var. 
flava is from the holotype, the only specimen known. 
The distribution of P. beringensis (closed square) is 
based on the dot map in Probatova (1985), P. byr- 
rangensis (diagonal hatching) is based on the dot map 
in Malyshev and Peshkova (1990), and P. jenisseiensis 
(horizontal hatching) is based on a description of the 
distribution in Tzvelev (1964). 


Results 
A morphological and habitat description of CAN 
535850 is given in Appendix 2. Figure 1 shows the 


distribution of CAN 535850, labelled “Egg River,” 
compared with all the taxa in Puccinellia sect. Pseudo- 
colpodium. 

Using Anderson’s Flora of Alaska and Adjacent Parts 
of Canada (Welsh 1974), CAN 535850 keyed out to 
Puccinellia wrightii (as Colpodium wrightii Scribn. & 
Merr.). In Grasses of the Soviet Union (Tzvelev 1976), 
it keyed out intermediate between P. colpodioides (as 
P. wrightii subsp. colpodioides (Tzvelev) Tzvelev) and 
P. byrrangensis. 

Single characters 

Table | presents a morphological comparison of the 
Puccinellia taxa under study. The values of CAN 
535850 and the values that overlap with those of CAN 
535850 are highlighted in bold. Figure 2 shows pho- 
tographs of inflorescences and spikelets comparing 
CAN 535850 with the other closely related species. 

CAN 535850 had 14-18 similarities with taxa in P. 
section Pseudocolpodium (except for P. wrightii var. 
flava) and 10-12 similarities to taxa in the P. arctica 
aggregate (Table 1, column SIM). The first nine char- 
acters in Table | show the overlap between CAN 


502 


535850 and species in P. section Pseudocolpodium 
that are different, for the most part, than the values in 
the P. arctica aggregate. One character in Table 1, ratio 
of inflorescence length to plant height, showed no 
overlap of CAN 535850 with any members of the P. 
section Pseudocolpodium, but instead showed overlap 
with members of the P. arctica aggregate. 

Within P. sect. Pseudocolpodium, the taxon with the 
most similarities to CAN 535850 was P. wrightii var. 
wrightii, overlapping in 18 characters (Table 1). The 
next closest were P. colpodioides (14) and P. vahliana 
(14). The characters that were similar to P. wrightii 
var. wrightii, but not to P. colpodioides or P. vahliana, 
are number of branches at lowest node of inflores- 
cence, first glume width, and palea hair. 

Inflorescences of CAN 535850 (Figure 2a), P. 
wrightii (Figure 2c), and P. poacea (Figure 2e) were 
typically open, except when young and then closed 
and contracted. The inflorescence of P. vahliana (Fig- 
ure 2b) was typically contracted, although sometimes 
slightly more loosely than in the photograph. The 
inflorescence of P. colpodioides (Figure 2d) was vari- 
able in shape, most often being contracted as in P. 
vahliana, but sometimes slightly more open as in Fig- 
ure 2d and more closely resembling CAN 535850. 


Multivariate analysis 

The PCA (Figure 3a) shows that on Factor | (influ- 
enced by lemma, palea, and glume lengths, and extent 
of hair on paleas and lemmas) and Factor 2 (influ- 
enced by first glume width and anther length) species 
in P. section Pseudocolpodium separate from the P. 
arctica aggregate, and CAN 535850 groups in the cen- 
tre of P. section Pseudocolpodium. When the P. arctica 
aggregate was removed (Figure 3b), CAN 535850 
OTUs grouped between P. colpodioides and P. 
wrightii on Factor 1, which was most influenced by 
measurements of the inflorescence shape, and clusters 
more with P. wrightii on Factor 2, which is more influ- 
enced highly negatively by the first and second glume 
widths and callus hair length, and highly positively by 
the number of branches at the lowest node. The OTUs 
representing the single collection from CAN 535850 
were dispersed almost as much as the separate collec- 
tions from each of the other taxa. 


Discussion 
Sectional alignment 

The specimen CAN 535850 aligns with Puccinellia 
sect. Pseudocolpodium based on the results from Table 
1 and the ordinations in Figure 3. The specimen CAN 
535850 and all of the species in P. sect. Pseudo- 
colpodium have generally glabrous glume and lemma 
margins, whereas those in the P. arctica aggregate (P. 
poacea shown in Figure 2) are minutely scabrous. In 
addition, CAN 535850 and most of P. section Pseudo- 
colpodium differ from the P. arctica aggregate by hav- 
ing generally longer glumes and longer lemmas with a 
wider hyaline margin. 


THE CANADIAN FIELD-NATURALIST 


Vol Tt 


Puccinellia section Pseudocolpodium: species 
alignment 

The only taxon from this section recorded to date 
from Canada is P. vahliana. CAN 535850 differs from 
P. vahliana by having diverging inflorescence branch- 
es, a smaller inflorescence to height ratio, narrower 
glumes, and by lacking thick villous hair at the base of 
the palea. CAN 535850 also has longer primary inflo- 
rescence branches, larger anthers, and a larger ratio of 
anther to lemma, but there is some overlap in these three 
characters. The wide glumes of P. vahliana account for 
a “wrapping” effect of the glumes around the lower lem- 
mas, especially visible when the spikelets are closed 
(see also P. colpodioides below). CAN 535850 is similar 
to P. vahliana in glume and lemma lengths (and ratios) 
and lemma hair characters. 

The next closest taxon geographically, P. wrightii 
var. wrightii, is from Alaska and easternmost Russia. 
In comparison with P. wrightii, CAN 535850 differs 
by being shorter, having smaller ligules and shorter 
lemmas and paleas. They share, however, similar 
inflorescence branch divergence, length and width of 
glumes, hyaline lemma margins, longitudinal folds of 
the lemmas, long anthers, and hairiness of florets. The 
specimen CAN 535850 generally has four or five 
branches at the lowest node of the inflorescence. 
While the original description of P. wrightii indicated 
the lowermost branches in pairs, specimens from Alas- 
ka had up to six branches at the lowest node. The CAN 
535850 specimen appears to be similar to P. wrightii, 
but less robust overall. This may be a consequence of 
our plant having grown in a more extreme environment. 

The description of P. wrightii var. flava is based on 
only one collection from Port Clarence, Alaska. This 
variety was originally described as only differing from 
the typical variety by having yellow florets. In addi- 
tion to this difference, we found that the lengths of first 
glume, second glume and lemmas are much larger 
than those of P. wrightii var. wrightii, with no overlap. 
The ratios of these values, however, are similar to the 
rest of P. sect. Pseudocolpodium. These very large 
length values account for the low number of similari- 
ties with CAN 535850. Moreover, CAN 535850 has 
straw-coloured to greenish-red florets with a bronze 
tinge, as opposed to yellow florets. 

The remaining taxa in this subgenus are known only 
from Russia. CAN 535850 keys out intermediate to P. 
colpodioides and P. byrrangensis in Tzvelev’s treat- 
ment (1976). Puccinellia colpodioides is a smaller 
plant than P. wrightii, and the lemma length of CAN 
535850 overlaps more with the smaller lemma of P. 
colpodioides than with that of P. wrightii. Some of the 
inflorescence characters (primary branch length, maxi- 
mum divergence of inflorescence branches) of CAN 
535850 are similar to those of P. colpodioides, but the 
former differs by having a smaller ratio of inflores- 
cence to plant height, a generally open inflorescence, 
with 4-5 branches at the lowest node, narrow glumes, 


2005 CONSAUL, GILLESPIE, AND MACINNES: ALKALI GRASS FROM THE ARCTIC ARCHIPELAGO = 503 


-RANTLEM 
AHYAL 


LEMV 


LEMV 


‘EGLW * PALH 


10) 


FIGURE 2. Photographs of five species in Puccinellia section Pseudocolpodium taken under a stereomicroscope. (a—e) Inflores- 
cences, scale bar length = 1 cm. (a) CAN 535850, (b) Puccinellia vahliana, (c) P. wrightii, (d) P. colpodioides, (e) P. 
poacea. (f-}) Spikelets, scale bar length = 1 mm. (f) CAN 535850, (g) Puccinellia vahliana, (h) P. wrightii, (i) P. colpo- 
dioides, (j) P. poacea. Black and white arrows show features that differ from those of CAN 535850. Abbreviations for 
the character codes are given in Table 1. 


504 


FACTOR(2) 


2 -1 0 1 2 


3 FACTOR(1) 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


FACTOR(2) 


-2 -1 O- 1 a 3 
b FACTOR(1) 


FIGURE 3. Principal components | ee (a) Analysis on full dataset (b) Analysis on dataset without P. arctica aggregate. Sec- 
tion Pseudocolpodium: E = CAN 535850, V = P. vahliana, C = P. colpodioides, W = P. wrightii; Species of P. arctica 
aggregate, section unassigned: A = P. arctica, P. = P. poacea, R = P. agrostidea. 


and silky hairs on the palea keels rather than villous 
hairs on the lower palea keels, similar points of differ- 
ence to P. vahliana. Thus, although the specimen does 
resemble P. colpodioides superficially, close examina- 
tion of the spikelets suggests that CAN 535850 is not 
as Closely aligned to P. colpodioides as to P. wrightii. 
Moreover, P. colpodioides has wide glumes similar to 
those of P. vahliana, which makes these two species 
closer in appearance to each other than to CAN 
535850. 

Both CAN 535850 and P. byrrangensis have very 
wide hyaline margins, but this is a rather variable 
character in CAN 535850. These species also share 
obvious folds on the lemma. On the other hand, the 
panicle was described as contracted for P. byrrangen- 
sis, unlike CAN 535850 which is most often open. The 
palea keels are glabrous or at most very slightly pilose 
in P. byrrangensis, unlike those of CAN 535850 which 
are slightly scabrous distally. Thus, differences in 
inflorescence characters indicate that CAN 535850 is 
unlikely to be P. byrrangensis, a species endemic to 
the Taimyr, west Siberia and considered by Tzvelev 
(1964) to be intermediate between P. vahliana and P. 
colpodioides. 

The distribution gap between CAN 535850 and 


both P. beringensis and P. jenesseinsis make it unlikely » 


that the former belongs to either of these species. Puc- 
cinellia beringensis is reported to have paleas that are 
glabrous and smooth or with solitary spinules, and 
anthers 1-1.4 mm long (Tzvelev 1976). In contrast, 
CAN 535850 has paleas that are slightly scabrous and 
anthers 1.6-1.8 mm long. Moreover, Probatova (1985) 


did not even classify P. beringensis in section Pseudo- 
colpodium [although Tzvelev (1976) did], but instead 
placed it in section Puccinellia. Puccinellia jenessein- 
sis has paleas that are glabrous or with only a few hairs 
basally, and inflorescences that are lax and broadly dif- 
fuse (Tzvelev 1964, 1976). Tzvelev (1964) described 
the lemmas as definitely lacking longitudinal folds, 
whereas in 1976 he said the lemmas were often longi- 
tudinally folded when dry. CAN 535850 has slightly 
scabrous veins to the palea as well as a few basal hairs, 
inflorescences that are open at the base only, and 
strong longitudinal folds. 

From the above comparisons, CAN 535850 aligns 
most closely with typical P. wrightii, although the flo- 
rets were smaller overall, which may explain why it 
keyed out closer to the smaller taxon P. colpodioides. 
In summary, CAN 535850 had palea hair characters 
resembling P. wrightii, glume length similar to P. 
wrightii, lemma size characteristics of P. byrrangensis, 
P. colpodioides, and P. vahliana, and the hyaline mar- 
gins and longitudinal folds of P wrightii and P. 
byrrangensis. Although intermediate, it is unlikely a 
hybrid because it is allopatric from all taxa except P. 
vahliana. 

Although CAN 535850 is different from all of the 
other species of Puccinellia section Pseudocolpodium 
we do not choose to base the description of a putative 
new taxon on a single specimen. The species in section 
Pseudocolpodium are very similar morphologically 
and in our PCA form a continuum as was found for 
other Puccinellia species complexes in Consaul and 
Gillespie (2001). CAN 535850 falls with P. colpodi- 


2005 


oides between P. vahliana and P. wrightii in morpho- 
logical characteristics. It conservatively aligns closest 
to P. wrightii, to which it conforms to a small specimen, 
as much as to P. wrightii var. flava is larger. Puccinellia 
wrightii may be an aggregate species complex which 
includes the larger P. wrightii forms and the smaller 
forms such as P. colpodioides and CAN 535850. The 
latter two may provide an evolutionary link between 
the allopatric species P. wrightii and P. vahliana. 


Taxonomy 

Puccinellia wrightii var. flava (Scribn. & Merr.) 
Consaul, comb. nov. Based on Colpodium wrightii 
subsp. flavum Scribn. & Merr., Contributions of the 
United States National Herbarium 13: 75. 1910. The 
single collection upon which this name is based is more 
yellow in colour and has larger glumes and florets than 
in the typical variety. The collection locality lies within 
the distribution of P. wrightii var. wrightii and collection 
notes on specimens suggest the habitats are similar; we, 
therefore, give this taxon only varietal status. 
Note: The combination Puccinellia wrightii (Scribn. & 
Merr.) Tzvelev was published in two places in 1964: 
Fl. Arct. URSS (Arkticheskaya flora SSSR), Fasc. II, 
193. 1964 (IK), and Novosti Sist. Vyssh. Rast. 1964: 
19 (GCI). Notes in the respective publications indicate 
that the former was signed off for publication 18 July 
1964 and the latter 14 September 1964. Therefore, the 
former is the proper citation for the publication of the 
name P. wrightil. 


Acknowledgments 

We thank Carolyn Parker and Vladim Federoy for 
translation of Russian on herbarium labels, and we 
also thank the former for providing us with geo- 
graphical coordinates for some of the more obscure 
Russian localities. The curators at ALA and US are 
thanked for loans of specimens. We thank W. J. Cody 
for collecting P. arctica from the Yukon. The Polar 
Continental Shelf Project is thanked for funding and 
logistic support for field work, and the Aurora Re- 
search Institute and Nunavut Research Institute are 
thanked for logistic support in the field. J. McNeill, 
Royal Botanical Gardens, Edinburgh, is thanked for 
comments and help with nomenclature. Two anony- 
mous reviewers are thanked for constructive comments 
on the manuscript. This research was funded by the 
Canadian Museum of Nature. This is Polar Continental 
Shelf publication #023-06. 


Documents Cited (marked * in text) 

Aiken, S. G., L. L. Consaul, and M. J. Dallwitz. 22 
November 2000. Poaceae of the Canadian Arctic Archi- 
pelago: Descriptions, Illustrations, Identification, and 
Information Retrieval. Version: 22" November 2000. http: 
//www.mun.ca/biology/delta/arcticf/(1995 onwards). 


CONSAUL, GILLESPIE, AND MACINNES: ALKALI GRASS FROM THE ARCTIC ARCHIPELAGO 


505 


Watson, L., and M. J. Dallwitz. 1999. Grass Genera of the 
World: Descriptions, Illustrations, Identification, and Infor- 
mation Retrieval; including Synonyms, Morphology, Anato- 
my, Physiology, Phytochemistry, Cytology, Classification, 
Pathogens, World and Local Distribution, and References. 
Version: 18" August 1999. http://biodiversity.uno.edu/delta/ 
(1992 onwards). 


Literature Cited 

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Research Council, Research Press, Ottawa, Ontario, Can- 
ada. 

Consaul, L. L., and L. J. Gillespie. 2001. A re-evaluation of 
species limits in Canadian Arctic Island Puccinellia 
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Davis, J. I. 1983. A systematic treatment of the Puccinellia 
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Davis, J. I. 1993. Puccinellia howellii (Poaceae) is genetical- 
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Fernald, M. L., and C. A. Weatherby. 1916. The genus 
Puccinellia in eastern North America. Rhodora 18: 1—23. 

Gould, F. W., and R. B. Shaw. 1983. Grass Systematics, 
second edition. Texas A&M University Press, College Sta- 
tion, Texas. 

Hultén, E. 1968. Flora of Alaska and neighboring territories. 
Stanford University Press, Stanford, California. 

Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 
1990. Index herbariorum, Part I: The Herbaria of the 
World. 8" Edition. New York Botanical Garden, New York. 

Malyshey, L. I., and G. A. Peshkova. Editors. 1990. Flora of 
Siberia [Flora Sibiriae]. Volume 2 Poaceae (Gramineae). 
Novosibirsk. [In Russian]. 

McRae, D., R. Stitt, and N. C. Wilson. 1994. A southern 
breeding range extension of the Lesser Snow Goose, Chen 
caerulescens caerulescens, James Bay, Ontario. Canadian 
Field-Naturalist 108: 223. 

Polunin, N. 1940. Botany of the Canadian Eastern Arctic. 
Part. 1. Pteridophyta and Spermatophyta. National Museum 
of Canada Bulletin Number 92. 

Polunin, N. 1959. Circumpolar Arctic Flora. Clarendon 
Press, Oxford, United Kingdom. 

Probatova, N.S. 1985. Puccinellia. Pages 310-321 in Plan- 
tae vasculares orientis extremi Sovietici. Edited by S. S. 
Charkevicz, Tomus 1, Nauka, Leningrad. [In Russian]. 

Tzvelev, N. N. 1964. Puccinellia Parl. — alkali grass. In Flora 
of the Russian Arctic. Arkitcheskaya flora SSSR, Volume 
2 (Gramineae). Edited by A. I. Tolmachev. USSR Academy 
of Sciences, Komarov Botanical Institute. [Translated from 
Russian by G.C.D. Griffiths. Edited by J.G. Packer. Uni- 
versity of Alberta Press, Edmonton. 1995. pp. 237—263.] 

Tzvelev, N. N. 1976. Zlaki SSSR. Nauka, Leningrad, Russia. 
[Grasses of the Soviet Union, Translated from Russian 
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Welsh, S. L. 1974.. Anderson’s Flora of Alaska and adjacent 
parts of Canada. Brigham Young University Press, Provo, 
Utah. 


Received 22 August 2002 
Accepted 13 September 2005 


506 


Appendix 1 
Specimens examined in the morphological analyses (names 
of authors of current paper abbreviated by initials). Herbari- 
um code in parentheses; abbreviations from Holmgren et al. 
1990). Specimens used in PCA coded by 

CAN 535850 - N.W.T.: Banks Island, Egg River, Area 
#16, K. L. MacInnes s.n., 24 July 1971, (CAN). P. agrostidea 
— NUNAVUT, Banks Island: De Salis Bay. 71°26'N, 121°40'W., 
31 July 1949, A. E. Porsild 17614 (CAN) Holotype?; Ellesmere 
Island: Lake Hazen. 81°49'N, 71°20'W., 8-10 Aug. 1961, C. 
R. Harington, 355 (CAN)?; Caledonian Bay. 79°57.23'N, 
81°11.96"W., 9 Aug? 1972, Waterton, I. W207 (CAN)?; Vic- 
toria Island: Cambridge Bay. 69°07'N, 105°03'W., 12 August 
1959, Porsild, A. E. 21599 (CAN)?; 69°07'N, 105°03'W., 12 
August 1959, Porsild, A. E. 21597 (CAN)?. P. arctica — 
NUNAVUT: Victoria Island: Cambridge Bay, 69°08'N, 
LOS" 10 W,. LIG & LEC 6321 (CAN): LIG & ELC 6322, 
LJIG & LLC 6342 (CAN); NWT: Anderson River Delta, 
69°40.66'N, 128°54.92'W, LJG & LLC 6362 (CAN); Atkinson 
Point, appr. 70°N, 131°20'W, A. E. & R. T: Porsild 2549 
(CAN); Cape Dalhousie, 70°11.28'N, 129°40.96'W, A. E. & 
R. T:. Porsild 2710 (CAN); Tuktoyaktuk, 69°27'N, 
133°02'W, A. E. Porsild 7404°; J. M. Gillett 18669 (CAN)?; 
J. M. Gillett 18787 (CAN); Yukon: Herschel Island, W. J. 
Cody 36154 (CAN). P. colpodioides — RUSSIA: prov. Mag- 
adan, Wrangel Is., 27 July 1970, V. Petrovsky 6303 (ALA)?; 
R.P. Nebrobikuit, 28 July 1972 (ALA)?; Somnitelnay Bay, 
vic. Zvzdnyy, approx. 70°50'N, 179°30'W, V.V. Petrovsky. 7 
August 1971 (voucher 2n=14) (ALA)?. P. poacea —- NUNA- 
VUT: Axel Heiberg Island: Diana Lake, appr. 79°30'N, 
88°30'W, A. E. Porsild 18640 (CAN)?; A. E. Porsild 18641 
(CAN)?; Mokka Fiord, 79°29'N, 87°22'W, LJG & C. Vogel 
6087 (CAN); Ellesmere Island: Hazen Camp, 81°49'N, 
71°20'W, LIG & C. Vogel 6215 (CAN); LJG & C. Vogel 6216 
(CAN); LJG & C. Vogel 6217 (CAN); Hot Weather Creek, 
79°58'N, 84°26'W, LIG & C. Vogel 6132 (CAN); LJG & C. 
Vogel 6144 (CAN); LJG & C. Vogel 6150 (CAN); LJG & C. 


Appendix 2 


Morphology and Habitat Description of CAN 535850 

Plants 8-18 cm tall; uppermost ligule 0.9-1.0 mm long; 
inflorescence 2.0-4.0 cm long, inflorescence length to plant 
height ratio 0.17-0.25; inflorescence lowest branches ascend- 
ing to horizontal, (2-) 4-5 branches at the lowest node; first 
glume 2.1-2.2 mm long, 0.6-0.8 mm wide; second glume 
2.9-3.2 mm long, 0.7-1.0 mm wide; lemma 3.5-4.0 mm long, 
with moderate to very wide hyaline margin, longitudinal ridges 
distinct, base pilose to moderately villous on and between 
the veins; palea keels distally moderately scabrous, basally 
glabrous to slightly pilose; anthers 1.6-1.8 mm long. Differs 
from P. colpodioides and P. vahliana by its open panicle, 
smaller ratio of inflorescence to height, relatively narrower 
glumes, and non-villous palea keel bases; from P. wrightii by 
its larger ratio of inflorescence to height and shorter lemma 
with a wider hyaline margin; from P. wrightii var. flava by its 
non-yellow colour of florets and smaller glumes, lemma and 
palea, and longitudinal ridges on lemma. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Vogel 6151 (CAN); Slidre Fiord, appr. 80°00'N, 85°57'W, J. 
S. Tener 33 (CAN); J.S. Tener 35, (CAN) Holotype?; J. S. 
Tener 34 (CAN)P?; Tanquary Fiord, 81°24'N, 76°52'W, J. S. 
J. Haight 6 (CAN)?. P. vahliana — NUNAVUT: Baffin 
Island: Nanisivik Airport, 73°02'N, 84°33'W, LJG, LLC & 
R. J. Soreng 6691 (CAN); Savage Harbour. 61°50'N, 
65°45'W, V. C. Wynne-Edwards 7301 (CAN)?; Cornwallis 
Island: Resolute Bay, 74°41'N, 94°50'W, LJG & C. Vogel 
6249 (CAN); A. E. Porsild 21649 (CAN)?; Devon Island: 
Dundas Harbour, 74°31.3'N, 82°33.5'W, LJG, LLC & R. J. 
Soreng 6697 (CAN); Ellesmere Island: Caledonian Bay, 
79°57.23'N, 81°11.96'W, LLC & LJG 2219 (CAN); Craig 
Harbour, 76°12'N, 81°O1'W, J. D. Soper 111374 (CAN)?; 
Hazen Camp, 81°49'N, 71°20'W, LJG 6012 (CAN); NWT, 
Banks Island: near Cape Lambton, 71°05'N, 123°09'W, A. E. 
Porsild 17539 (CAN)?; Melville Island: Ibbett Bay Camp, 
75°54'N, 114°30'W, S. G. Aiken & S. A. Edlund 3936 
(CAN)?. P. wrightii - RUSSIA: Chukotka Peninsula, in the 
vicinity of the Chaplin Cape, 20 July 1958, R. Satpuisau s.n. 
(ALA)?; Chukotka Peninsula, SE tip of Cape Chaplino, 
approx. 64°25'N, 172°15'W, 20 July 1958, V. Gavreluk 
(ALA); Chukotka Peninsula, eastern region, S coast of Lav- 
rentiya Bay, vicinity of Lavrentiya, approx. 65°45'N, 
171°W, 29 August 1971, N. A. Sekretarova, A. K. Sitinn, B. 
A. Yurtsev (ALA)?; Chukotka Peninsula, eastern tip, vicinity 
of Uelen, approx. 66°10'N, 169°50'W, 25 Aug. 1971, N. A. 
Sekretarova, A. K. Sitinn, & B. A. Yurtsev (ALA)?; SW por- 
tion of Chukotka peninsula, right bank of Senevem R., 15 
km above mouth. 24 July 1983, A. E. Katenen & N. A. 
Sekretarova (ALA); W. Chukotka, Anyuyskiy Mountains, 
upper Vernitakatvem River, 12 July 1974, 7: Koraseva and 
V. Petrovsky (ALA), voucher 2n=14; Siberia, Arakam- 
tchetchene or Kayne Island, C. Wright (US ex NA 592344) 
(Holotype). USA: Alaska, Teller Quad., Cape Prince of 
Wales, 65°37'N, 168°05'W, 18 August 1982, 7. Kelso 82- 
230 (ALA); P. wrightii var. flava: Alaska, Port Clarence, 
65°16'N, 166°51'W. F) A. Walpole 189la (US 379007) 
(Holotype). 


Specimen CAN 535850 was found above the Egg River 
floodplain in a transition zone between a slightly steeper slope 
with soil stripes and more pronounced Dryas/Salix hummocks 
below, and a more open summit zone with Dryas, Salix, Sax- 
ifraga tricuspidata Rothb. and Cetraria above. The collection 
site had a gradual slope, with 30% ground cover of rounded 
soil mounds or micropolygons, and predominantly Dryas/Salix 
cover. The plant was found on the bare to partly bare tops of 
these low Dryas mounds, with salt crust, sand, pebbles and 
desiccation cracks containing lichens and mosses. Scattered 
small depressions in the area contained predominantly sedges 
and grass, as well as Cassiope tetragona (L.) D.Don. Other 
associated herbs in the general area were: Hulteniella integri- 


folia (Richardson) Tzvelev, Parrya arctica R.Br., Saxifraga 


oppositifolia L., Astragalus alpinus L., Pedicularis capitata 
Adams, and Pedicularis langsdorfii subsp. arctica (R. Br.) 
Pennell. 


‘Prairie Grouse’, 7ympanuchus cupido X phasianellus, Hybridization 
on Manitoulin Island, Ontario 


H. G. LUMSDEN 
144 Hillview Road., Aurora, Ontario, L4G 2M5 Canada 


Lumsden, H. G. 2005. “Prairie Grouse”, Tympanuchus cupido xX phasianellus, hybridization on Manitoulin Island, Ontario. 
Canadian Field-Naturalist 119(4): 507-514. 


Greater Prairie Chickens started their range expansion from Wisconsin about 1900. They reached Sault Ste. Marie, Ontario, 
by 1925 and completed colonization of Manitoulin Island by 1945. In the fall of 1932, an irruption of Northern Sharp-tailed 
Grouse from the Hudson Bay Lowlands occurred. Residents of Manitoulin distinguished the two species and reported the 
presence of “square-tails” and “‘sharp-tails” in the winter of 1932-1933. There is no specimen evidence or verbal reports that 
Northern Sharp-tailed Grouse reached Michigan during that irruption. Prairie Sharp-tailed Grouse were in the northern 
peninsula of Michigan by 1930. In 1941, Michigan Department of Natural Resources made a planting of 37 Prairie Sharp- 
tailed Grouse on Drummond Island. Thence they spread east and the first lek was found at the west end of Manitoulin Island 
in 1952. By 1960, these birds had virtually colonized the whole island. Development of a hybrid index from the morphology 
of specimens of skins and skeletons from Manitoulin indicated that more than 50% of all birds in the 1960s were hybrids. In 
Michigan, less than 1% of the birds necropsied or checked in hunters’ bags were hybrid. It seems likely that ethological isolation 
broke down on Manitoulin Island. The booming display of the Prairie Chicken and the tail rattling display of the Sharp-tailed 
Grouse can be broken down into their component parts. They appear to be homologous to five discrete displays of the Spruce 
Grouse, grouped in different patterns in each of the two lek species. There was little hybridization between Prairie Sharp- 
tailed Grouse and Prairie Chickens that had lived sympatrically for thousands of years. Northern Sharp-tailed Grouse had 
probably never lived sympatrically with Prairie Chickens and the evolution of the perception of species distinctions may not 
have evolved to the point where hybridization was restricted to a rare event. 


Key Words: Greater Prairie Chicken, Tympanuchus cupido, Sharp-tailed Grouse, Tympanuchus phasianellus, hybrids, behav- 


iour, Manitoulin Island Ontario, Michigan. 


In Ontario during the last 150 years the ranges of 
many species of birds have changed dramatically (Sny- 
der 1957), mostly as a result of introduction and later 
intensification of European farming methods and log- 
ging. Their effect in Michigan and the adjacent areas 
of Ontario have altered the habitat and allowed the 
sequential invasion of Manitoulin Island by Greater- 
Prairie Chickens (Tympanuchus cupido) and two races 
of Sharp-tailed Grouse (Tympanuchus phasianellus). 

Land use in the Manitoulin area is largely dictated 
by the geology of the Island. Almost horizontally bed- 
ded ordovician limestone slightly tilted toward the south 
is generally covered with shallow soils. Only about 
12% of the island has soils deep enough for farming. 
Over 40 500 ha (about 25% of the island) is limestone 
plain which is largely open because of grazing by cat- 
tle and pulp-wood cutting. Both of these uses created 
and have steadily enlarged the prairie-like appearance 
of the landscape (Braffette and Brown 1948*). How- 
ever, that land use is changing today and there is less 
grazing and pulp-wood cutting. 

Although unlike the traditional range occupied by 
Prairie Chickens and Sharp-tailed Grouse prior to Euro- 
pean settlement, it proved to be suitable and was col- 
onized. In this paper I outline the history of the three 
invasions of Manitoulin Island by three forms of “Prai- 
rie Grouse” and offer some hypotheses concerning the 
development of the hybrid population. Although not 


an inhabitant of prairie, I shall refer to the Northern 
Sharp-tailed Grouse (7: p. phasianellus) and the mixed 
population of Manitoulin Island collectively as “Prairie 
Grouse” in this paper. 


History of Prairie Grouse on Manitoulin 
Island 

Prairie Chickens and Sharp-tailed Grouse were sym- 
patric over a wide area in southern Wisconsin in pre-and 
early settlement times (Schorger 1944). They were 
ecologically isolated, with the Prairie Chicken occupy- 
ing the open Long-Grass Prairie and the Sharp-tailed 
Grouse confined to the Oak openings and brushy areas. 
With the advent of lumbering and settlement, new range 
appeared to the north, east and west of their historic 
distribution and extraordinary range expansions took 
place. Prairie Chickens moved west almost to the foot- 
hills of the Rocky Mountains and as far as one hundred 
and fifty kilometres north of Edmonton (Houston 
2002). They also spread to the northeast, followed 
twenty years later by the Prairie Sharp-tailed Grouse 
(T. p. campestris), terminating on Manitoulin Island, 
Ontario. 

Between 1875 and 1920 the original range of both 
species was being converted to farmland and new range 
in the north was being created by lumbering followed 
by fire (Hamerstrom et al. 1957). Parts of every county 
but not all townships in Wisconsin were affected. 


507 


508 


Prairie Chickens 

The first Prairie Chickens to enter the Upper Penin- 
sula of Michigan presumably came from Wisconsin 
about 1900 (Ammann 1950*, 1957). They spread across 
the Peninsula reaching the western half of Chippawa 
County in 1923-1924. This county includes the north- 
eastern part of the Upper Peninsula adjacent to St. 
Joseph’s Island and the Sault Ste. Marie area of On- 
tario. 

Baillie (1947) wrote, “The first indication that these 
birds were expanding their range eastward from Michi- 
gan came from the west side of St. Joseph’s Island, 
south of Sault Ste. Marie, where a few appeared about 
1925. R. H. Burns of Sault Ste. Marie assured us that 
by 1946 the birds, although not plentiful, had moved 
over onto the Canadian mainland and were in occu- 
pation of a narrow belt of cleared land along the St. 
Mary’s River up to a distance of three miles inland 
where the edge of the Pre-Cambrian Shield prevents 
their further progress.” W. E. Gimbly (Ontario Depart- 
ment of Lands and Forests, now Ministry of Natural 
Resources, personal communication) saw some Prairie 
Chickens at Echo Bay, twelve miles east of Sault Ste. 
Marie in the late 1930s and W. St. John (L & F per- 
sonal communication) said that they also reached the 
Kirkwood area north of Thessalon about that time. 
This invasion of this part of Algoma including Mani- 
toulin Island by Prairie Chickens occurred a few years 
before the spectacular irruption of Northern Sharp- 
tailed Grouse of 1932-1933. 

Some of the residents of Manitoulin Island still 
remembered in the 1950s the 1932 invasion of Sharp- 
tailed Grouse. John and Cecil Merrylees saw a flock of 
“Prairie Grouse” feeding on grain spilled from sleighs 
on Christmas Eve 1932, on the road west of Indian 
Point Bridge in Burpee Township. “The flock consisted 
mostly of “sharptails” but some “square” tails were 
included” (Baillie 1947). This was the first eyewitness 
account of both Prairie Chickens and Sharp-tailed 
Grouse occurring together on the Island. The Indian 
Point bridge is about 56 kilometres from the western 
end of the Island. It is unlikely that the Prairie Chickens 
reached this point during the first year of their presence 
on the Island. They had been advancing at an average 
rate of 13.5 kilometres per year in Michigan. Ammann 
(1950*) reported that they reached the lower Seney 
marshes in 1910-12 about 270 kilometres to the west. 
If their rate of advance was relatively steady they may 
have arrived on Manitoulin [sland about four years 
earlier, about 1928. By 1932 there were probably sev- 
eral leks with breeding populations established on suit- 
able range that lies between Meldrum Bay and Indian 
Point. . 

Baillie (1947*) also recorded five additional obser- 
vations in the 1930s. Only one, from Larry Donaldson, 
refers definitely to “square” tails being seen near Gore 
Bay in Gordon Township about 1937. By 1939 they 
had apparently reached Great Cloche Island near Lit- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


tle Current in Howland Township where W. Wilkinson 
reported seeing six to eight. They had moved about 
55 km in seven years or at an average rate of eight 
kilometres per year. By 1942, they were reported on 
Ten Mile Point in Sheguiandah Township and at Man- 
itowaning in Assiginack Townships by C. J. Young 
(Ontario Lands and Forests, personal communication). 
They had virtually colonized the entire island. 

The only specimen of a Prairie Grouse preserved 
from the early invasion of the north shore of Lake 
Huron east of Sault Ste. Marie was collected by Pro- 
fessor N. Rae Brown in the Kirkwood Forest Manage- 
ment Unit north of Thessalon on 13 September 1942. 
It is a hybrid Prairie Chicken x Sharp-tailed Grouse. 
Figure | maps the advance of the Prairie Chicken in 
the Michigan-Manitoulin area. 


Northern Sharp-tailed Grouse 

The breeding distribution of the Northern Sharp- 
tailed Grouse in Ontario, based largely on the reported 
location of leks, is confined to peatland muskegs and 
burns in the north. The First Nations people report 
leks within a few km of the Hudson Bay coast. Along 
the southern fringe of their breeding range they breed 
regularly south to the northern Canadian National Rail- 
way line. In some areas they breed further south to 
where better drained land-forms permit development 
of forest within which there are a few outlying mus- 
kegs large enough to support breeding populations; 
for example, near Matheson, Tionaga, and the Black 
Sturgeon area near Lake Nipigon. 

There are areas of abandoned or little used cultivated 
land that lie adjacent or close to occupied muskegs 
but the Northern Sharp-tailed Grouse seems to be un- 
able to colonize them. Their inability to adapt to grass- 
land habitat perhaps explains their failure to persist 
in areas to which they irrupted in 1932-1933. 

There are about 260 0007 km of muskeg country in 
northern Ontario which forms the range of the North- 
ern Sharp-tailed Grouse. In the northern part of this 
range there are annual fall migrations of these birds into 
the river courses and to the south particularly in the 
rockier western parts of Ontario. 

Every few years some birds move into the Cochrane 
area where they come to the gravel roads for grit and 
are easily shot. When populations are exceptionally 
high, these fall migrations take the form of irruptions 
that carry the birds far to the south. Snyder (1935) sug- 
gests an irruption in 1865-1866. Fleming (1906) doc- 
uments an irruption in 1896-1897 that carried birds 
as far south as Muskoka and Parry Sound Districts. 

Snyder (1935) described in detail the irruption of 
Northern Sharp-tailed Grouse that started in mid-Octo- 
ber 1932. By March 1933 the birds had penetrated to 
many localities in the Parry Sound District. A speci- 
men was preserved which was taken as far south as 
Bracebridge in Muskoka District, but Snyder did not 


“mention that they reached Manitoulin Island. 


2005 


SAULT 
STE. MARIE 
s 


UPPER PENINSULA 
OF MICHIGAN 


. 
1910 


1923 
SS) @ Mackinaw City 


jz 


Cc eboygan 


LUMSDEN: GROUSE HYBRIDIZATION ON MANITOULIN ISLAND 


509 


19 
DRUMMOND 
E COCKBURN 


Stig 


FiGuRE |. Approximate speed of advance of Prairie Chickens in northern Michigan and Manitoulin Island, Ontario. 


Although no specimens of Northern Sharp-tailed 
Grouse from the Island were preserved from the 1932- 
1933 irruption, there is confirmation in the surviving 
files of the Department of Game and Fisheries! that 
they did reach Manitoulin Island in 1932. In a letter 
dated 20 February 1933 J. M. Parks, Superintendent 
at North Bay, writing of Sharp-tailed Grouse, “On 
November 5" last it was estimated that several hundreds 
of birds alighted in the town of Elk Lake, and at the 
present time small flocks are to be found in the vicinity 
of North Bay, Sudbury and the Manitoulin Island.” He 
wrote a second letter on 6 March 1933, “Flocks have 
again been reported in the vicinity of Webbwood and 
the Manitoulin Island.” During this irruption Northern 
Sharp-tailed Grouse spread over an area of approxi- 
mately 100 000 square kilometres to the south of their 
normal breeding range in Ontario. That did not include 
the area south of the French and Mattawa Rivers where 
only a few flocks appeared, nor did it include the area 
the birds occupied in Quebec. 

In the Sault Ste. Marie area, a flock of Sharp-tailed 
Grouse appeared near the school on Lot 11 Concession 
A of St. Joseph’s Township during the winter of 1932 
according to Murray Smith (Lands and Forests, per- 
sonal communication). His brother, F. B. H. Smith, saw 
a flock of “Prairie Grouse” (species uncertain) about a 
hundred strong, in the same place in 1935. 

Snyder (1935) listed seven observations of young 
birds or nests with eggs reported from south of the 


normal breeding area in the summer of 1933. Although 
there may be an error in any one of these reports, Sny- 
der pointed out, that the aggregate provides “un- 
questionable evidence that the species nested in terri- 
tory occupied by the immigrants”. However, by the 
summer of 1934 they had virtually disappeared from 
their irruptive range. 


Prairie Sharp-tailed Grouse 

The eastward spread of the Prairie Sharp-tailed 
Grouse occurred about twenty years after that of the 
Prairie Chicken. They were assisted in their movement 
by a release made by the Michigan Department of Con- 
servation. Ammann (1947*) wrote of the Upper Penin- 
sula: “By the early thirties Sharptails had spread to 
and become abundant in most suitable areas in the 
western third of the peninsula. Except for Drummond 
Island, however, they were not commonly noted in 
the extreme eastern end (Eastern Chippawa and Mac- 
kinac Counties) until 1949.” Thirty-seven banded 
Sharp-tailed Grouse were trapped, moved and released 
in February and March 1941 near Johnswood on 
Drummond Island (Ammann 1947*). They undoubt- 
edly bred that spring. In 1942 only one bird of fifteen 
shot was found to have been banded. The population 
resulting from this successful planting was probably 
responsible for the subsequent invasion of Cockburn 
and Manitoulin Islands. Four specimens preserved by 
H. McQuarrie and B. Smith were shot on Cockburn 


' Amalgamated in 1946 with the Department of Lands and Forests, the name was later changed to the Ministry of Natural 


Resources (M.N.R) 


510 


Island in November 1950. The birds were abundant 
there and had evidently been present for some years. 

The first specimen of a Prairie Sharp-tailed Grouse 
on Manitoulin Island was a female shot by W. Ritching 
on the east bluff at Gore Bay in Gordon Township in 
1948. This specimen, given to the author without a 
label, is now in the Royal Ontario Museum. It is a fe- 
male in extremely worn plumage. Searches and inten- 
sive observations from blinds were carried out between 
1949 and 1952 on known leks. No Sharptails were dis- 
covered until 1952; it seems likely that this female 
penetrated well ahead of the main invasion. 

On 26 April 1952, I found a Sharp-tailed Grouse 
lek four kilometres west of Meldrum Bay in Dawson 
Township, Manitoulin. On 28 April 1952, there were 
fifteen birds present, four of which were females. Two 
birds were collected: one was a Sharp-tailed Grouse, 
and the second was a hybrid. Prairie Sharp-tailed 
Grouse probably had established their first leks on 
Manitoulin Island about 1950. On 25 April 1952, a 
single Sharp-tailed Grouse hen appeared at the lek at 
the airport near Gore Bay about 50 km from the west 
end of the Island. H. McQuarrie collected the first male 
Sharp-tailed Grouse recorded on the central part of the 
Island on the lek at Britainville, Campbell Township 
in 1953. Probably about this time males first arrived at 
the airport at Gore Bay. By 1960, Sharp-tail-like birds 
were abundant at the Gore Bay airport, Britainville, 
and in Billings Township where six specimens were 
collected on 24 September 1960. 

In the spring of 1962 the first Sharp-tailed Grouse 
appeared at the east end of Manitoulin Island in She- 
guiandah Township. John Budd (Lands and Forests, 
personal communication) found a dead bird that may 
have hit telephone wires beside the road but the species 
was still scarce. Only Prairie Chickens and hybrids ap- 
peared on the two leks in Sheguiandah Township that 
were studied in 1962. Figure 2 maps the advance of 
Prairie Sharp-tailed Grouse from Michigan to Mani- 
toulin Island. 

The Prairie Chicken stock had colonized, from St. 
Joseph’s Island to eastern Manitoulin, a distance of 
187 km in 20 years, an average of about nine km per 
year. It took the Prairie Sharp-tailed Grouse 21 years 
to spread from Drummond Island in their year of 
release to the eastern end of Manitoulin Island, a dis- 
tance of 137 kilometres at an average speed of six km 
per year. They were substantially slower than the Prairie 
Chickens twenty years earlier. 

The Hamerstroms (1951), summarizing Wisconsin 
recovery data, concluded, “...we are convinced... that 
Prairie Chickens really are somewhat more mobile than 
Sharptails most of the year.” It would appear that if 
fidelity to the home lek or other behavioural factors 
did not interfere with changes in home range, Prairie 
Chickens could be expected to colonize new areas at 
a faster rate than Sharp-tailed Grouse. Another factor 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


must be considered. Prairie Sharp-tailed Grouse have 
replaced Prairie Chickens in most of Northern Michi- 
gan and the adjacent parts of Ontario. This would sug- 
gest that there is competition between the two species 
in this kind of range. It is likely that the Prairie Chick- 
en was only able to colonize the Sault Ste. Marie- 
Manitoulin part of Ontario in the absence of competi- 
tion from Prairie Sharp-tailed Grouse. The mechanisms 
of competition are at present unknown. It is possible 
that Sharp-tailed Grouse did not colonize this new 
range at the same rate as they might have done had 
the range been vacant of competitors. 


History of Prairie Grouse in Northern 
Michigan 

The same genetic stocks of Prairie Chickens and 
Prairie Sharp-tailed Grouse over-ran the Upper Penin- 
sula of Michigan a few years before they reached Man- 
itoulin Island but there seems to be no evidence that 
the northern race of the Sharp-tailed Grouse reached 
Michigan during its irruption in 1932-1933. 

I have used Ammann’s (1950*) unpublished report 
on the spread of both species in Michigan as a basis, 
with modifications according to his (personal commu- 
nication) subsequent suggestions to estimate the years 
during which both species were present at the same 
time in each county. 

Using only those years, Table 1 summarizes the data 
resulting from necropsies by game division biologists 
and checked in hunters’ bags in the field in the Cusino 
area. Ammann (personal communication) suggested that 
some hybrids could have been missed before staff were 
aware that hybrids might be present. In the necropsy 
sample, 6 of 420 (1.4%) of the birds checked were 
identified as hybrids. In the Cusino field checks, only 
4 of 1206 (0.3%) were identified as hybrids. Combin- 
ing these hunter bag checks in fall with observations 
of living birds on their leks in spring, sixteen positive 
and eight probable hybrids were identified. There were 
three additional hybrid specimens shot in fall and one 
seen in spring on Drummond Island. 

Johnsgard and Wood (1968) reported many hybrids 
from three provinces and seven states where the two 
species occur sympatrically. Their report indicates 
widespread casual hybridization but nowhere was a 
hybrid population described. It is clear that the hybrids 
recorded in Northern Michigan were similarly products 
of casual cross matings and that at no time did a hybrid 
population similar to that of Manitoulin Island develop. 

We have two areas of range, both of which were oc- 
cupied by the same genetic stock of Wisconsin Prairie 
Chickens, but those areas were initially invaded by a 
different stock of Sharp-tailed Grouse. Casual hybri- 
dization occurred in Northern Michigan with the Prairie 
Sharp-tailed Grouse, whereas a hybrid population re- 
sulted on Manitoulin Island with the Northern Sharp- 


tailed Grouse. 


2005 


UPPER PENINSULA 


OF 


MICHIGAN 
OSeney 


O Cusino 


S rs} Mackinaw City ge 


Cheboygan 
UJ 


LUMSDEN: GROUSE HYBRIDIZATION ON MANITOULIN ISLAND 


1941 
Planting 


S11 


1951 
DRUMMOND 
' s 
COCKBURN 


ie '. Meidrum Ba 


Blind River 


Little 
Current 


FIGURE 2. Approximate speed of advance of Sharp-tailed Grouse in northern Michigan and Manitoulin Island, Ontario. 


Observations on Leks 

Starting in 1951, watching from a blind on selected 
leks on Manitoulin Island, I made judgements on wheth- 
er each male was a hybrid, a Prairie Chicken, undeter- 
mined or a Sharp-tailed Grouse. I used their plumage 
markings, aberrations in their displays and the colour 
of their booming sacks and toes. By 1951, Prairie Sharp- 
tailed Grouse had only reached the extreme western 
end of Manitoulin Island and none had appeared on 
the four leks intensively studied. Counts of these leks 
(Table 2) suggested that of 48 males present, 28 (58%) 
were hybrids, 16 (33%) were Prairie Chickens, 4 (8%) 
were undetermined, and none were Sharp-tailed Grouse. 
Those in the undetermined category had the appear- 
ance of Prairie Chickens but slight shortening of the 
pinnae, or occasional peculiarities of behaviour caused 
me to question their identity. 

By 1962, Prairie Sharp-tailed Grouse had taken over 
at Gore Bay Airport. There were only 3 (23%) hybrids, 
no Prairie Chickens, but 10 (7%) apparently pure Prai- 
rie Sharp-tailed Grouse. 

In 1961, on the extreme eastern end of the Island at 
the Sheguiandah lek, there were still 10 (83%) hybrids, 
2 (17%) Prairie Chickens, and no Sharp-tailed Grouse. 
Much the same ratios prevailed at Sheguiandah in 


1962 when 6 (85%) were hybrids and there was only 
1 (15%) Prairie Chicken. However in 1964, Prairie 
Sharp-tailed Grouse had appeared at Sheguiandah and 
in 1966 at Ten Mile Point. The proportions of hybrids 
in each year for each lek studied ranged from 23% to 
85% (Table 2). 


Discussion 

The hybrid frequency on Manitoulin Island amounts 
to more than 50% of the population (Lumsden in 
preparation) but in northern Michigan far less than 1%. 
There are thus significantly different rates of hybridi- 
zation in these two areas. Prairie Chickens and Sharp- 
tailed Grouse do not produce hybrid populations wher- 
ever they have been sympatric. Mayr (1942) outlined 
four categories of isolating mechanisms, one or more 
of which can operate to maintain the genetic integrity 
of a population. They are: (1.) ecological isolation; (2.) 
ethological isolation; (3.) mechanical isolation; (4.) 
genetic isolation. 

The hybrid population that developed on Manitoulin 
Island indicates that there could be no genetic, mechan- 
ical or ecologically isolating mechanisms operating. 
It is likely that ethological isolating mechanisms pro- 
vided the most effective barrier between the species else- 


TABLE |. Number of Prairie Chickens, Sharp-tailed Grouse and hybrids from the upper peninsula of Michigan necropsied by 
Game Division biologists 1935-1957 and checked by field staff at Cusino 1939-1954. 


Prairie Chickens 


1935-1957 necropsies 49 
1939-1954 field checks 110 


Sharp-tailed Grouse Hybrids 
S71 6! 


1096 4? 


' first hybrid identified in 1939 
* two hybrids identified by G. A. Ammann 


S12 


TABLE 2. Number of male hybrids, Prairie Chickens, Undetermined, and Prairie Sharp-tailed Grouse tallied on various leks 


THE CANADIAN FIELD-NATURALIST 


in 1951 and 1952 and 1964-1966. 


Lek Prairie Sharp-tailed 
1951 Hybrid Chicken Undetermined Grouse 
Gore Bay Airport 10 (55%) 4 (22%) 4 (22%) 0 
Billing’s Township 2 (33%) 4 (66%) 0 0 
Britainville 6 (75%) 2 (25%) 0 0 
Tustians Farm 10 (63%) 6 (37%) 0 0 
Total 28 (58%) 16 (33%) 4 (8%) 0 
1952 

Gore Bay Airport 6 (42%) 4 (29%) 4 (29%) 0 
Sheguiandah 6 (85%) 1 (14%) 0 0 
Total 12 (57%) 5 (24%) 4 (19%) 0 
1964 

Sheguiandah 8 (67%) 1 (8%) 0 3 (25%) 
1965 

Sheguiandah 6 (60%) 1 (10%) 0) 3 (30%) 
1966 

Sheguiandah 4 (50%) 2 (25%) 0 2 (25%) 
10 Mile Point 4 (57%) 0 0 3 (43%) 
Total 8 (53%) 2 (13%) 0 5 (33%) 


Vol. 119 


where and that this broke down on Manitoulin Island. 

In non-lek species, such as the Spruce Grouse (Can- 
achites canadensis) males that display in isolation on 
their territories usually must respond at one time to no 
more than a single visitor of their own species. Thus, 
on the approach of a rival male the territory holder 
could react with one or more displays. On the approach 
of a female the male might respond with courtship 
displays (Lumsden 1961) and finally by copulation if 
the hen were receptive. 

In a lek-species, however, where territories are very 
small and many males may be displaying in close prox- 
imity, a single male may have to respond simultaneous- 
ly to a challenge from a dominant neighbour, to an inva- 
sion by a subdominant rival or to a visit by a receptive 
female. Each lek species has a complex display which 
seems to be appropriate in a variety of circumstances 
and which is performed endlessly. The lek display of 
the Prairie Chicken and the tail-rattling display of the 
Sharp-tailed Grouse are the multi-purpose displays 
which function in these complex social situations. 

Schenkel (1956 and 1958) proposed that these mul- 
tiple purpose displays of the above species are homo- 
logous with the tidbitting displays of many species of 
pheasant. I cannot agree with this. First, the tidbitting 
displays so far described for pheasants are all frontal 
displays, while the booming and tail-rattling displays 
are lateral in orientation. Second, one can hardly find 
a single component element in common between tid- 
bitting and the grouse displays. 

If we break down these multipurpose displays of lek 
species into their component parts and look for homo- 
logies in the displays of a solitary species (e.g., the 
Spruce Grouse), we find elements from four different 
Spruce Grouse displays — hooting, tail flick, strutting 


and head and tail down displays are telescoped into 
the booming display of the Prairie Chicken. In the case 
of the Sharp-tailed Grouse, tail-rattling display elements 
from the tail swishing, strutting, head-jerk, and head 
and tail down displays appear. Table 3 summarizes how 
these elements are grouped. It is important to note that 
the multipurpose displays of Prairie Chickens and 
Sharp-tailed Grouse, while homologous with several 
displays of a solitary species, have the component ele- 
ments so grouped that the two displays present totally 
different sounds and appearances to an observer. These 
differences constitute the behavioural isolating mecha- 
nism. Anyone who is familiar with the two species will 
recognize that it is possible to identify the lek, whether 
Prairie Chicken or Sharp-tailed Grouse, as far away as 
the birds can be heard, perhaps two kilometres. Sexu- 
ally active hens would have no difficulty in locating 
the lek of their own species in an area of sympatry. 
Similarly, should a hen make a mistake and visit a lek 
of the wrong species, the courtship behaviour directed 
at her would normally tend to correct her mistake. 

If these species-specific displays constitute the main 
isolating mechanism, it is likely that perception by 
the female of the differences must be present to make 
them effective. I suggest that the Prairie race of the 
Sharp-tailed Grouse (Tp. campestris) that may have 
lived sympatrically with Prairie Chickens for thousands 
of years, through selection, produced a biotype of fe- 
male in which perception was very sharp. Females of 
the muskeg-dwelling Northern Sharp-tailed Grouse (7p. 
phasianellus), which may never have lived in sympatry 
with the Prairie Chicken, were not subjected to this 
kind of selection. This may explain the breakdown of 
species isolation on Manitoulin Island and the pro- 
duction of a hybrid population. 


2005 


LUMSDEN: GROUSE HYBRIDIZATION ON MANITOULIN ISLAND 


5s, 


TABLE 3. Elements in Spruce Grouse displays grouped in the booming and tail-rattling displays of Prairie Chickens and 


Sharp-tailed Grouse. 


Spruce Grouse 
Booming 


Hooting Display 
inflation of oesophagus X 
hooting vocalization x 
Tail Flick 
(inhibited locomotion) 
Rapid steps 
Tail Flicked open 
Wings lowered 
Strutting/Tail Swishing 
Spreading alternate sides of tail 
Wings lowered X 
Inflation of oesophagus X 
Head-Jerk or Squatting 
Wings held open and away 

from the body 
Head and Tail Down 
Head and neck extended forward xX 


xx xX 


There are further factors that may have contributed 
to the production of hybrids. Snyder (1935) reported 
that the sex ratio of Northern Sharp-tailed Grouse that 
irrupted in 1932 was grossly distorted. Of the 65 speci- 
mens that were preserved in museums, the sex ratio 
was 74% females to 26% males, most of which were 
in their first year. Furthermore, the birds that reached 
Manitoulin Island had no traditional leks at which they 
could seek matings in the spring of 1933. It is apparent 
that leks of the resident Prairie Chickens in western 
Manitoulin attracted traditionless Northern Sharp-tailed 
Grouse. This occurred again when Prairie Sharp-tailed 
Grouse invaded Manitoulin Island in the early 1950s. 
They did not immediately establish leks of their own but 
appeared at the existing Prairie Chicken and hybrid leks. 

An additional factor may have enhanced the op- 
portunity for hybridization and perhaps reduced the 
influence of the male Sharp-tailed Grouse on the 1933 
year class. A lek is composed of a group of males in 
which an hierarchy of dominance prevails. Repeated 
observations on both Sharp-tailed Grouse and Prairie 
Chicken leks have confirmed that only very few males 
perform all the matings. In some cases, a single alpha 
male may copulate with 75% of the females that visit 
the lek (Lumsden 1965*). Among Prairie Chickens, 
Robel (1967, 1970) reported that two males performed 
89% of 121 copulations. The extent to which the 
Sharp-tailed Grouse males contributed to the 1933 year 
class would depend on their ability to achieve high 
status in the lek hierarchies. This would be difficult for 
them since most were less than one year old. 

Snyder (1935) wrote that the Northern Sharp-tailed 
Grouse disappeared from their acquired range very 
rapidly and that by the winter of 1933-1934 there were 


Prairie Chicken 


Nuptial Bow 


Sharp-tailed Grouse 


Tail Rattling Cooing Nuptial Bow 


Seldom XxX 
Seldom XxX 


very few left. This population from the muskeg is 
apparently not adapted to life in grassland and it is 
likely that most disappeared from Manitoulin within 
a year. Some may have returned to the north and others 
died. The hybrids produced in 1933 and the Prairie 
Chickens were left to continue to interbreed and to 
colonize the rest of the Island 

Table 2 shows that during the invasion by Prairie 
Sharp-tailed Grouse (7p. campestris), no males had 
reached the Gore Bay Airport by 1951. However, by 
1962 77% of the 13 males present were Sharp-tailed 
Grouse. At the same time at Sheguiandah on the ex- 
treme east end of the island in 1962, there were no 
Sharp-tailed Grouse among the seven males present. 
By 1964, 25% of 12 males tallied there were Sharp- 
tailed Grouse. By 1966, the proportion of Sharp-tailed 
Grouse had risen to 33% at Sheguiandah and 10 Mile 
Point. 

Field studies were not continued after 1970 but nat- 
uralists have maintained an interest in the “Prairie” 
Grouse of Manitoulin since. Observations made by 
Jerry Guild (Ontario Field Ornithologists News 1997) 
on the Gore Bay Airport on 15 April, 1997 yielded a 
count of 50 Sharp-tailed Grouse. Jean Iron (personal 
communication) recorded 75-100 Sharp-tailed Grouse 
on 15 April 2000; 30 on 12 April 2003; and 60 on 11 
April 2004 at the Gore Bay Airport. No Prairie Chick- 
ens or hybrids were recorded in these years. No natu- 
ralist seems to have checked the “Prairie” Grouse on 
the eastern end of the island. In view of the success 
of Sharp-tailed Grouse in the Gore Bay area and the 
time lapse since the last checks, it seems likely that 
Sharp-tailed Grouse have now displaced Prairie Chick- 
ens and hybrids throughout the island. 


514 


Acknowledgments 

I thank G. A. Ammann for his generous loan of 
unpublished reports. I thank F. N. and F. Hamerstrom 
for fruitful discussions on the biology of Prairie Grouse 
and loan of specimens. The land owners on Manitoulin 
Island generously permitted me to work on their prop- 
erty. I thank W. Riching, H. McQuarrie, and B. Smith 
for the many specimens they donated and for help with 
fieldwork. I am most grateful to A. J. Erskine and an 
unknown referee who greatly improved this paper. 


Documents Cited (marked * in text) 

Ammann, G. A. 1947. History and status of Sharp-Tail Grouse 
and Prairie Chickens on Drummond Island. Unpublished 
Report. Michigan Department of Conservation. 

Ammann, G. A. 1950. History of the Prairie Grouse in Mich- 
igan. Unpublished Report 1119 Michigan: Department of 
Conservation. 

Baillie, J. L. 1947. Report on the Prairie Chicken Project 
Manitoulin 1947. Unpublished Report. Ontario Department 
of Lands and Forests. 

Braffette, R. W., and D. K. Brown. 1948. Survey of Mani- 
toulin Island. Unpublished report. Ontario Department of 
Lands and Forests. 

Lumsden, H. G. 1965. Displays of the Sharp-tailed Grouse. 
Unpublished Research Report Number 66 Technical Series. 
Ontario Department of Lands and Forests. 


Literature Cited 

Ammann, G. A. 1957. The Prairie Grouse of Michigan. Mich- 
igan: Game Division, Department of Conservation. 200 
pages. 

Fleming, J. H. 1906. Range of the Sharp-tailed Grouse in 
eastern Canada. Ontario Natural Science Bulletin 2: 19. 
Hamerstrom Jr., F. N., and Frances Hamerstrom. 1951. 

Mobility of the Sharp-tailed Grouse in relation to its ecol- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


ogy and distribution. American Midland Naturalist 46: 
174-226. 

Hamerstrom Jr., F. M., O. E. Mattson, and Frances Ham- 
erstrom. 1957. A guide to Prairie Chicken management. 
Wisconsin Conservation Department. Technical Wildlife 
Bulletin 15: 127 pages. 

Houston, C. S. 2002. Spread and disappearance of the Greater 
Prairie Chicken (Tympanuchus cupido) on the Canadian 
Prairies and adjacent areas. Canadian Field-Naturalist 116: 
1-21. 

Johnsgard, P. A., and R. E. Wood. 1968. Distributional 
changes and interaction between Prairie Chickens and 
Sharp-tailed Grouse in the Midwest. Wilson Bulletin 80: 
173-188. 

Lumsden, H. G. 1961. Displays of the Spruce Grouse. Cana- 
dian Field-Naturalist 75: 152-160. 

Mayr, E. 1942. Systematics and the origin of species. Colum- 
bia, Ohio. Columbia University Press. 1-334. 

Robel, R. J. 1967. Significance of booming grounds of Great- 
er Prairie Chickens. Proceedings of the American Philo- 
sophical Society 111: 109-114. 

Robel, R. J. 1970. Possible role of behaviour in regulating 
Greater Prairie Chicken Populations. Journal of Wildlife 
Management 34: 306-312. 

Schenkel, Von R. 1956 and1958. Zur Deutung der Balzleis- 
tungen einiger Phasianiden and Tetraoniden. Der Ornithol- 
ogische Beobachter 53: 182-201; 55: 65-95. 

Schorger, A. W. 1944. The Prairie Chicken and Sharp-tailed 
Grouse in early Wisconsin. Transactions of the Wisconsin 
Academy of Science Arts and Letters 35: 1-59. 

Snyder, L. L. 1935. A Study of the Sharp-tailed Grouse. Biol- 
ogy Series. University of Toronto Press: 1-66. 

Snyder, L. L. 1957. Changes in the avifauna of Ontario. 
Pages 26-42 in Changes in the fauna of Ontario. Edited 
by F. A. Urquhart. University of Toronto Press. Toronto. 


Received 25 June 2004 
Accepted 29 March 2005 


Plumage and Internal Morphology of the “Prairie Grouse”, 
Tympanuchus cupido X phasianellus, of Manitoulin Island, Ontario 


HARRY G. LUMSDEN 
144 Hillview Road, Aurora, Ontario L4G 2M5 Canada 


Lumsden, Harry G. 2005. Plumage and internal morphology of the “Prairie Grouse”, Tympanuchus cupido X phasianellus, of 
Manitoulin Island, Ontario. Canadian Field-Naturalist 119(4): 515-524. 


I made comparisons among populations of Greater Prairie-Chickens, Sharp-tailed Grouse and their hybrids on Manitoulin Island 
of pinnae length, breast, flank and tail feather patterns, tail graduations, wing chord, and skeletal characteristics. Estimates of 
the proportion of hybrids from these individual characters ranged from 32% to 60%. Lek behaviour of hybrids was a mixture 
of the patterns of the parent species. The colour of the booming sacks varied and displayed the range between the parent 
species.There was a tendency with time for the characters of Prairie Chickens to decline coincident with an increase in Sharp- 
tailed Grouse characters. Both Prairie Chickens and Sharp-tailed Grouse are lek species in which an alpha male may consum- 
mate most of the the matings. The speed with which morphological change took place in Manitoulin Island is to be expected 
where a non-random mating system of this kind prevails. 


Key Words: Greater Prairie-Chickens, Tympanuchus cupido pinnatus, Sharp-tailed Grouse, Tympanuchus phasianellus, proportion 


of hybrids, morphology, behaviour, population changes, Heath Hen, 7ympanuchus cupido cupido, species relationships. 


The occupation of Manitoulin Island, Ontario, by 
Greater Prairie-Chickens (Tympanuchus cupido pinna- 
tus) and their hybrids with Northern Sharp-tailed 
Grouse (T. p. phasianellus) was essentially complete 
by 1945 and offers clues to species relationships. Trap- 
ping of these birds started in the winter of 1948-1949 
and field studies continued intermittently until 1966, 
with hunting season collections continuing until 1970. 
Between 1943 and 1946, Prairie Sharp-tailed Grouse 
(Tympanuchus p. campestris) moved into the western 
end of the island and reached the eastern end about 
1960 (Lumsden 2005S). 

Specimens of hybrids showed the full range of char- 
acters between Greater Prairie-Chickens and Sharp- 
tailed Grouse. The purpose of this study is to estimate 
the proportion of hybrids present in the Manitoulin 
Island population, its change with time and to describe 
its morphology. Manitoulin specimens are curated at 
the Royal Ontario Museum in Toronto. 


Methods 

Manitoulin specimens were sorted into three groups. 
The samples taken from 1949 to 1959 consisted of 
Greater Prairie Chickens (Tympanuchus cupido pin- 
natus) and their hybrids with Northern Sharp-tailed 
Grouse (Tympanuchus phasianellus phasianellus). They 
included specimens from the western end of the island 
and later the eastern end near or on leks where no 
Prairie Sharp-tailed Grouse (Tympanuchus p. campes- 
tris) to that time had been recorded. 

From 1960 to 1963, Prairie Chickens, hybrids and 
Prairie Sharp-tailed Grouse were present in all parts of 
the island. Between 1964 and 1970, Prairie Chickens 
had disappeared and the earlier hybrids were inter- 
breeding with Prairie Sharp-tailed Grouse. For compar- 


ison with the Manitoulin specimens, Prairie Chickens 
were borrowed from five museums with specimens from 
Manitoba, Saskatchewan, Alberta, Michigan, Wisconsin, 
Minnesota, Iowa, Montana, North and South Dakota, 
Nebraska, [Illinois and Oklahoma. All the Northern 
Sharp-tailed Grouse were collected in northern Ontario. 


Morphological Characters 

Pinnae, breast, flank and tail feathers were collected 
from trapped birds that were released elsewhere. On 
the pinnae feathers from 9 to 11 mm of the proximal 
end of the shaft were white and were embedded in tis- 
sue. Measurements were taken from the terminus of 
the white on the shaft to the tip of the feather. Measure- 
ments were taken from study skins by sliding a thin 
ruler under the longest feather and reading from the 
skin to the tip. 

The pinnae lengths were divided into five groups: 
Class 1: 84-72 mm; Class 2: 71-59 mm; Class 3: 58-46 
mm; Class 4: 45-33 mm; and Class 5: 32-20 mm 
(Table 1). Feathers from the upper breast (Figure 1) 
(Table 2) and the lower flanks (Figure 2) (Table 3) 
were selected to represent the range of pattern variation 
seen in the three populations. These were arranged into 
five pattern classes ranging from the barred Prairie 
Chicken pattern (Class 1) to the acute- angled arrow- 
shaped pattern (Class 5) of the Sharp-tailed Grouse. The 
equivalent feathers on all specimens were matched 
with these patterns and a relevant value assigned. Fig- 
ure 3 illustrates the steeply graduated tail of a male 
Sharp-tailed Grouse. Graduation was the length of the 
outer tail feathers subtracted from the length of the cen- 
tral tail feathers. Measurements were taken from where 
the feather emerged from the skin to the tip. 

Division of the measurements of the degree of gradu- 
ation of the tail into five groups were Class 1: 30-44 mm, 


515 


SLUG) 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE |. Classes of length of the pinnae of male Prairie Chickens, the Manitoulin population and Northern Sharp-tailed Grouse. 


Class 2 
Prairie Chicken 100 (77%) 30 (23%) 
Manitoulin 
1949-1959 7 (11%) 30 (39%) 
Northern Sharp-tailed Grouse 0 0 


3 4 5 Total 
0 0 0 130 
11 14%) 4 (5%) 24 (32%) 76 
0 0 19 (100%) 19 


Praire Chicken Hybrids Sharp-tailed Grouse 
No. 1857 No. 1346 No. 352 No. 1201 No. 1353 
e) e) © ‘) e) 
Index Value | 2 3 4 5) 


FiGuRE |. Upper breast feathers from a Prairie Chicken, three hybrids and a Sharp-tailed Grouse showing range of variation. 


Class 2: 45-58 mm, Class 3: 59-73 mm, Class 4: 74- 
87 mm and Class 5: 88-102 mm (Table 4). Figure 4 
illustrates the pattern on the tail feathers of a male 
Prairie Chicken, a female and four male Sharp-tailed 
Grouse. The pattern on the latter varies in males and is 
often an indicator of sex (Table 5). 

There was a small overlap in the length of the wing 
(chord) between Prairie Chickens and Northern Sharp- 
tailed Grouse. The wing lengths were divided into 
Class 1: 239-230 mm; Class 2: 229-220 mm; Class 3: 
219-210 mm; Class 4: 209-200 mm; and Class 5: 199- 
190 mm (Table 6). 


Skeletons were saved from as many specimens as 
possible. Measurements were taken from the tip of the 
ilium process to the ventral surface of the pubic bone. 
In cases where the ilium process projected below the 
pubic bone, a minus value was assigned. Figure 5 
shows the lateral view of the ilium and ischium of 
Prairie Chickens, Heath Hens (Tympanuchus cupido 
cupido), (see note page 000) and Northern Sharp-tailed 
Grouse. Size classes summarized in Table 7 were as 
follows: Class 1: -2.5- +0.7 mm; Class 2: 0.8-4.0 mm; 
Class 3: 4.1-7.3 mm; Class 4: 7.4-10.6 mm; and Class 
5: 10.7-13.9 mm. The vertebral column consists of a 


TABLE 2. Classes of patterns on the upper breast feathers of male Prairie Chickens, the Manitoulin Island population and North- 


ern Sharp-tailed Grouse. 


Class | D 
Prairie Chickens 175 (100%) 0 
Manitoulin 
1949-1959 14 (42%) 9 (27%) 
Manitoulin 
1960-1963 8 (12%) 8 (12%) 
Manitoulin 
1964-1970 2 (4%) 5 (9%) 


Northern Sharp-tailed Grouse 0 0 


3 4 5 Total 
0 0 0 175 
6 (18%) 3 (9%) kM (3%) a3 
12 (19%) 9 (14%) 27 (42%) 64 
It re) IW. (20%) 24 (45%) 53 
0 0 66 (100%) 66 


es Pe Seer 


2005 LUMSDEN: PLUMAGE AND INTERNAL MORPHOLOGY OF ““PRAIRE GROUSE” S17 


_ me 


Praire Chicken Hybrids Sharp-tailed Grouse 
No. 1857 Nos i171 No. 1202 No. 1199 No. 1353 
@) @) o) o) o) 
Index Value 1 ‘ 2 3 d 5 


FiGuRE 2. Lower flank feathers from a Prairie Chicken, three hybrids and a Sharp-tailed Grouse showing range of variation. 


TABLE 3. Classes of patterns on the feathers of the lower flank feathers in male Prairie Chickens, the Manitoulin Island pop- 
ulation and Sharp-tailed Grouse. 


Class 1 2 3 4 5 Total 
Prairie Chickens 174 (100%) 0 0 0 0 174 
Manitoulin 
1949-1959 11 (33%) fl 3%) Zz (6%) 26 %) 7. Q1%) 53 
Manitoulin 
1960-1963 5 (8%) 3 (4%) qT Ciioy oe a) 42 (66%) 64 
Manitoulin 
1964-1970 0 4 (7%) 3 (6%) 8 (15%) 38 (72%) 35 
Northern Sharp-tailed Grouse 0 0 0 3 (4%) 63 (95%) 66 


TABLE 4. Degree of graduation of the tail of male Prairie Chickens, the Manitoulin population and Northern Sharp-tailed Grouse. 


Class 1 2 3 4 5 Total 
Prairie Chickens 144 (99 %) Dire) 0 0 0 146 
Manitoulin 
1949-1959 39 (67 %) 13) (22%) 4 (7%) 2 (3%) 0 58 
Manitoulin 
1960-1963 5 (16%) 3 (9%) 13) (40%) 7 22%) 4 (13%) Se 
Manitoulin 
1964-1970 0 2 (ir%) 8 (42%) 9 (47%) 0 19 


| Northern Sharp-tailed Grouse 0 0 0 29" (69%) 45) (61%) 74° 


518 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE 5. Frequency of males from Manitoulin Island classified as Sharp-tailed Grouse by criteria other than the tail pattern 
and as hybrids, which showed a pattern of lines parallel to the shaft on the central tail feathers. 


Classified as Sharp-tailed Grouse 


With Pattern Without Pattern 
1949-1959 l 0 
1960-1963 4 13 
1964-1970 if 3 
Total 12 (43%) 16 (37%) 


Classified as Hybrids 
With Pattern Without Pattern 
0 39 
2 28 
3 21 
7 (Fe 88 (93%) 


TABLE 6. Classes of the length of the wing (chord) of male Prairie Chickens, the Manitoulin population and Northern Sharp- 


tailed Grouse. 


Class 1 2 
Prairie Chickens 31 (20%) 105 (69%) 
Manitoulin 
1949-1959 0 11 (36%) 
Manitoulin 
1960-1963 0 8 (13%) 
Manitoulin 
1964-1970 0 2 (4%) 
Northern Sharp-tailed Grouse 0 0 


3 - 5 Total 
ap a s/o) We 3) 0 153 
14 (47%) 5 (17%) 0 30 
37 (60%) 16 (26%) 1 @%) 62 
16 (31%) 22 (42%) 12). (23%) 52 
2 (2%)  S584073%) 20 (23%) 80 


TABLE 7. Distance from the tip of the ilium process to the ventral surface of the pubic bone in male Prairie Chickens, the Mani- 


toulin population, and Sharp-tailed Grouse. 


Class 2 
Prairie Chickens 27 (87%) 4 (13%) 
Manitoulin 
1949-1959 12 (36%) it @G3%) 
Manitoulin 
1960-1963 2 (4%) 5) (10%) 
Manitoulin 
1964-1970 0 0 
Northern Sharp-tailed Grouse 0 0 


variable number of bones, particularly among the 
hybrids. I follow Campbell and Lack (1985) in their 
definitions of subdivisions. The cervical or neck verte- 
brae are free with the posterior one carrying a floating 
rib. The thoracic or dorsal vertebrae carry ribs articu- 
lated through sternal ribs with the sternum. 

Four thoracic vertebrae are normally fused to one 
another but a fifth is free. The sixth is fused to the lum- 
bar vertebrae and carries a rib that articulates with the 
neighbouring sternal rib but not directly with the ster- 
num. The lumbar and sacral vertebrae, which form the 
synsacrum, are fused to one another but some to the 
ilium. I have counted as caudal or tail vertebrae those 
that are free. However, the number varies with age. 
The anterior one may fuse with the synsacrum in old 
birds. I did not include in Table 8 very young birds that 


3 4 J Total 
0 0 0 31 
hb QIe) 37 Ge) 0 33 
6 (12%) 9 (18%) = 28. (56%) 50 
5 OGY 234%) 920 (57%) SE) 
0 if (@8i%) ~~ 28° (72%) 39 


had not completed fusion of the synsacrum. The verte- 
bral column ends with the pygostyle. 


Results 
Length of the Pinnae 

Prairie Chickens have a tuft of feathers 70-85 mm 
long on the sides of the neck above the anterior edge of 
the booming sac. On the Sharp-tailed Grouse, the 
equivalent feathers are only about 23 mm long. Table 1 
presents the number of birds with pinnae in each class 
for Prairie Chickens, Manitoulin Island specimens, and 
Northern Sharp-tailed Grouse. Because 77% of the 
Prairie Chickens are in Class 1, we should consider that 
77% of 7 Manitoulin birds are also Prairie Chickens 


- leaving only two hybrids in that class. In Class 2, there 


are 30 (23%) of the Prairie Chickens. Therefore, 23% 


2005 


LUMSDEN: PLUMAGE AND INTERNAL MORPHOLOGY OF “‘PRAIRE GROUSE” 


FIGURE 3. Feathers illustrating the steeply graduated tail of a male Sharp-tailed Grouse. 


of 30 Manitoulin birds must be considered to be Prairie 
Chickens, leaving 23 hybrids. All the Manitoulin birds 
in Class 3 and 4 must be considered hybrids, totalling 
15. All the Sharp-tailed Grouse were placed in Class 5 
SO we must consider the 24 Class 5 Manitoulin Grouse 
as Sharp-tailed Grouse. Left are 40 probable hybrids 
out of 76 specimens from Manitoulin or 52 %. It was 
unfortunate that nearly all specimens collected in fall 
during hunting seasons of 1960 to 1970 were molting 
with pinnae and tail feathers not fully grown. 

Fred and Frances Hamerstrom graciously let me 
measure their collection of 275 pinnae of Prairie 
Chickens collected in Wisconsin. The mean for those 
Wisconsin specimens was 77.7 mm whereas the maxi- 
mum for Manitoulin birds was 75 mm. 


Pattern on the Upper Breast Feathers 

As all Prairie Chickens were assigned to Class | 
(Table 2), we must assume that all Manitoulin speci- 
mens in Class | were also Prairie Chickens. All Sharp- 
tailed Grouse fell into Class 5, therefore all Manitoulin 
birds in Class 5 should be considered to be Sharp-tailed 


Grouse. The remaining 74 Manitoulin grouse constitut- 
ing 49% of the sample can be regarded as hybrids. 


Patterns on the Lower Flank Feathers 

All Prairie Chickens were placed in Class | (Table 
3), So we must consider all the Manitoulin specimens 
falling into this class as being Prairie Chickens. In 
Class 4 only 4% were Sharp-tailed Grouse leaving 16 
hybrids. With Class 5 birds from Manitoulin, all but 3 
(5%) must be considered as Sharp-tailed Grouse. There 
were 49 hybrids among the Manitoulin sample or 32%. 


Tail Graduation 

The tail feathers of a Sharp-tailed Grouse are very 
steeply graduated. The outer pair is only about 35% of 
the length of the central ones. In the Prairie Chicken, 
the tail is almost square with the outer rectrices meas- 
uring about 71% of the pair at the centre. The degree of 
graduation was the difference in length between the 
outer and central pair. It should be noted that the central 
pair of tail feathers in both species does not emerge 
from the tissue on the same plane as the eight pairs of 
rectrices. They grow on the same plane as the upper tail 


520 


THE CANADIAN FIELD-NATURALIST 


Voly ie 


TABLE 8. Counts of cervical, thoracic, synsacral and caudal vertebrae of Prairie Chickens,the Manitoulin populations and 


Sharp-tailed Grouse, all years and both sexes combined. 


Cervical Thoracic 

Vertebrae Vertebrae 
15 16 DY 6 a 
Greater Prairie Chicken 33 0 0 38 0 
Manitoulin Population 56 2 LL P28) 6 
Sharp-tailed Grouse a3 0 0 47 l 


“ one extra thoracic but missing one lumbar 


coverts and should properly be recognized as modified 
tail coverts. Table 4 presents the measurements broken 
down into five classes. As 44 of the Manitoulin birds 
were placed in Class | with 99% of the pure Prairie 
Chickens, there is a possibility that none of them were 
hybrids. Two (1%) of the Prairie Chickens were placed 
in Class 2. A few of the Class 2 Manitoulin birds 
may have been pure Prairie Chickens. Adjusting their 
numbers by 1% should provide an estimate of 17 
hybrids, 29 (39%) of the Sharp-tailed Grouse were 
placed in Class 4. The number of hybrids in Class 4 
must be reduced from 18 to 11. In Class 5, the num- 
ber of true hybrids is likely to have been 2. Most of the 
specimens in Class 5 were collected during the molt 
hence the small sample size. Out of 109 Manitoulin 
specimens, 55 or 50% do not conform to this charac- 
ter of either parent species and can be considered 
hybrids. 


Tail Pattern in Sharp-tailed Grouse 

The central tail feathers of many male Sharp-tailed 
Grouse (e.g., #217, Figure 4) are distinctive with lines 
running parallel to the shaft. Females have irregular 
bars running across the feather. A proportion of the 
males lack the line pattern and have a barred pattern 
that is somewhat similar to that of a female (Figure 
4). About 86% of the Prairie Sharp-tailed Grouse 
males have the distinctive line pattern; 14% do not 
(Snyder 1935; Manwiler 1939). In the male Northern 
Sharp-tailed Grouse I have examined, 12% are lack- 
ing the lines running parallel to the shaft. Examina- 
tion of the specimens from Manitoulin Island did not 
adhere to the above pattern. Of 28 males classified as 
Sharp-tailed Grouse by other criteria, one would expect 
that 3 (12%) or 4 (14%) would lack the line pattern. 
In fact, 57% were without this character, a rate four 
or five times higher than expected. Of males classi- 
fied as hybrids, 93% lacked the line pattern (Table 5). 

The dorsal surfaces of the rectrices of male Sharp- 
tailed Grouse are very pale grey and are white at the tips. 
In hybrids, this part of the tail is a darker grey even in 


Synsacrum Unfused Number of birds 
Caudale in the 
sample 
11 12F 13 2) 6 fi) 
Sie Sl" MS 1G Paar 38 
Zi Dit AG Khe 2A VI 58 
AN 46: 03 0 Sil 8 Bye 


those that approach Sharp-tailed Grouse in other charac- 
ters. Prairie Chickens have all the rectrices and even 
the central modified tail coverts dark in colour with 
some lighter coloured bars or spots. 


Length of the Wing 

The lengths of the wings of Prairie Chickens and 
Northern Sharp-tailed Grouse overlap slightly, but the 
former average distinctly larger (Table 6). There were 
no Manitoulin birds in Class 1. In Class 2, 69% were 
Prairie Chickens, leaving 7 probable hybrids. In Class 
3, of the 67 Manitoulin specimens, about 14% could 
be either Prairie Chickens or Northern Sharp-tailed 
Grouse, leaving 58 probable hybrids. Class 4 contained 
43 Manitoulin specimens but there were 58 (73%) 
Northern Sharp-tailed Grouse, leaving 12 probable 
hybrids. Class 5 contained 25% of the Northern Sharp- 
tailed Grouse, therefore 10 of 13 Manitoulin birds were 
most likely to be hybrids. Those adjustments produced 
an estimate of 60% hybrids. 


Distance from the Tip of the Ilium Process to the 
Pubic Bone 

There is a substantial difference in the shape of the 
pelvic area between the Prairie Chicken and the North- 
ern Sharp-tailed Grouse. In the former, the illum 
process projects ventrally so that it overhangs the ischi- 
um and occasionally even projects beyond and below 
the pubic bone (Figure 5). In the Northern Sharp-tailed 
Grouse it projects laterally, barely obscuring the 
ischiadic foramen. Table 7 records that 87% of the 
Prairie Chickens were in Class 1. We can assume that 
only 2 of the 14 Manitoulin birds in Class | were 
hybrids. In Class 2, 13% must be regarded as Prairie 
Chickens leaving 14 probably hybrids. All 16 Class 3 
Manitoulin birds were hybrids. Among the Sharp- 
tailed Grouse, 28% were in Class 4, so our estimate of 
hybrids is 17. As Class 5 contained 72% of the Sharp- 
tailed Grouse, we can estimate only 14 of 48 from 
Manitoulin as hybrids. Thus, 63 Manitoulin specimens 
were probable hybrids constituting 53% of the 118 
samples. 


2005 


Praire Chicken 
No. 1800 No. 1289 No. 114 
©) ) @ 
Index value | 2 


LUMSDEN: PLUMAGE AND INTERNAL MORPHOLOGY OF ““PRAIRE GROUSE” 


BPA) 


{| 
| 
| 


Sharp-tailed Grouse 


No. 1803 No. 2028 No. 217 
Co CO @ 
2 5 5 


FiGuRE 4. Central rectrices from a male Prairie Chicken, one female and four male Sharp-tailed Grouse showing variation in 


markings. 


Vertebral Column 

Skeletons available, not all complete, were 38 
Prairie Chickens, 58 from the Manitoulin population 
and 53 Northern Sharp-tailed Grouse. Table 8 summa- 
rizes the numbers of vertebrae in each sector of the 
spine for the three populations. The only variation in 
the cervical vertebrae occurs among the Manitoulin 
birds, 2 of which had an extra cervical vertebra. Counts 
of the thoracic vertebrae among the Manitoulin spec- 
imens revealed that one bird possessed only 5 but six 
had 7 (17% of the series). Among the caudal vertebrae 
of all three populations there was variation. None of 
the Prairie Chickens had seven, none of the Northern 
Sharp-tailed Grouse had five, but nineteen hybrids had 
five and one had seven. 


Not only is there variation in the number of verte- 
brae in each sector of the spine of the Manitoulin birds, 
but there is also variation in the structure of the bones. 
Table 9 summarizes six categories of aberrations that 
appeared in the vertebral columns. Of the 38 Prairie 
Chickens studied, three (8%) had aberrant structures. 
Of the 58 Manitoulin specimens, 24 (41%) were abnor- 
mal in structure. Among the Northern Sharp-tailed 
Grouse, two (4%) of 53 showed abnormalities. Many 
of those aberrations might not affect the functioning of 
the bird as far as survival was concerned. More serious 
abnormalities such as fusion or missing vertebrae 
might influence survival, possibly in situations of 
escape from avian predators. More serious but unmea- 
sured in this study would be possible aberrations in 


522 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLE 9. Number of specimens with aberrant structures in the vertebral column in Prairie Chickens, the Manitoulin population 


and Sharp-tailed Grouse. 
Category of aberrance 


Five dorsals fused l 
All six dorsals fused 0 
Vestigial rib on the anterior lumbar 0 
One missing lumbar 0 
Incomplete parapophyses connecting 


the lumbar with the ilium 2 
Posterior dorsal not properly fused to the lumbar 0 
Total 3 


muscle tracts, mixed behavioural signals or improper 
responses to external threats. 

In mature Prairie Chickens there are five or six free 
caudal vertebrae (Table 8). In Sharp-tailed Grouse 
there are six or seven. The evolution of the tail rattling 
display in the latter would likely have carried with it a 
change in the mobility and muscular structure of the 
tail, hence the increase in the length of the tail and its 
capacity for movement. The pygostyle is generally con- 
sidered to be composed of a number of fused caudal 
vertebrae. As both these species have eight pairs of rec- 
trices, it may be reasonable to assume that the pygostyle 
is composed of at least eight fused caudal vertebrae. 


Behaviour 

Schwartz (1945) gave a very complete description 
of the booming display of the Prairie Chicken. Typical- 
ly its component parts consist of raising the pinnae 
vertically above the head, rapid stomping of the feet 
while standing still or running forward, raising or 
lowering the head, inflating the oesophagus so that the 
orange “booming sacs” are distended; a rapid opening 
of the tail feathers, which produces an audible click, 
slight lowering of the wings but retaining the carpal 
joint within the covering flank feathers, and uttering 
three “booming” notes which sound like blowing over 
the open neck of a large bottle. This call may carry 
for several kilometres. The homologous tail rattling 
display of the Sharp-tailed Grouse consists of holding 
the horizontal body high off the ground on extended 
legs, lowering the head, raising the feathers dorsal to 
the cervical apteria, exposing the purple booming sacs, 
extending the wings from the body sometimes at full 
stretch with a downward curve of the primaries, erect- 
ing the tail at least vertically and often tilted slightly 
forward, spreading each side of the tail alternately at 
high speed making a loud rattling sound, and uttering 
a note that sounds like a cork being withdrawn from a 
bottle or a shrill “chilk” or hoarse “cha” call. 

A second display homologous with the “booming” 
segment of the Prairie Chicken display is called “‘coo- 
ing”. The bird stands still, extending the neck forward 
and slightly upward, inflating the oesophagus and 
hence distending the purple cervical apteria, bowing 


Prairie Chicken 


Manitoulin Population = Sharp-tailed Grouse 


2 0 
3 0 
9 l 
l 0 
8 | 
l 0 
24 z 


the neck slightly downward and uttering a loud cooing 
note, usually keeping the carpal joint of the wing cov- 
ered by the flank feathers (Lumsden1965*). 

Those hybrids that most nearly resembled Prairie 
Chickens produced the booming display, sometimes 
typically but frequently with three weak notes and 
some omitted the third note. One hybrid boomed with 
a single note, drooping his wings as he uttered and 
spread his tail with an audible hissing sound. 

Another hybrid cooed instead of booming: he held 
his wings slightly open but flicked them fully open, 
spreading his primaries while vibrating his tail without 
the volume of rattling in a pure Sharp-tailed Grouse. 
He produced an audible click with his tail at the end of 
this performance. Another hybrid, when cooing, uttered 
a double note. A Prairie Chicken-like hybrid made a 
rustling sound with his tail when booming and extend- 
ed his wings slightly, retracting them at the third boom- 
ing note. A Sharp-tailed Grouse-like hybrid produced 
three wheezy notes in his tail-rattling display. There 
was much confusing variation in the displays of 
hybrids and individuals were not as consistent in the 
component parts of their display as the two parent 
species. 


Colours of Booming Sacs and Feet 

The resonance of the booming and cooing calls is 
produced by inflation of the oesophagus that distends 
the coloured skin of the cervical apteria. In Prairie 
Chickens, these booming sacs, or timpani as they are 
sometimes called, are orange in colour with a narrow 
pink rim adjacent to the feathers. In Northern Sharp- 
tailed Grouse (T. p. phasianellus), the booming saks 
are smaller than in Prairie Chickens and are purple. At 
the height of the mating season they may be almost as 
dark as blueberries. Those of the Prairie Sharp-tailed 
Grouse (T. p. campestris) are much paler and are pink- 
ish-purple in colour. The timpani of hybrids are very 
variable in colour. Those in the middle of the cline 
between the species may have a muddy violet edge 
with one to three small dull orange patches. Others 
may have a reduced orange-yellow patch with an 
enlarged dusky pink rim. 


2005 


Prairie Chickens have dusky orange toes during the 
breeding season. In Sharp-tailed Grouse, they are grey. 
Hybrids varied from orange through yellowish orange, 
to yellowish-grey to grey coloured feet. All these 
Prairie Grouse seem to molt the pectinations of their 
toes in mid-April and the displaying males, toward the 
end of the display period, show considerable wear on 
their nails. 


Discussion 

The hybrid indices summarized in Tables 1-4, 6-7 
suggested that the proportion of hybrids in the Mani- 
toulin population was between 32% and 60%. Certain 
features of the pinnae and tail suggest that hybridiza- 
tion may have been more extensive than this. Unex- 
pected in the morphology of the Manitoulin Prairie 
Grouse was the relatively short pinnae of the popula- 
tion. That none reached the mean length of 77.7 mm of 
Wisconsin Prairie Chickens was a surprise. The longest 
measured 75 mm. Genetically, the Manitoulin birds are 
descended from the Wisconsin stock, 20 to 30 years 
before this study began and several hundred kilometres 
removed. This strongly suggests that hybridization on 
the island was more extensive than indicated by the 
other hybrid indices. The line pattern on the tail of 
Northern Sharp-tailed Grouse was missing in a propor- 
tion 4 or 5 times that expected of Sharp-tailed-type 
Manitoulin Grouse. Perhaps it is not unexpected that 
93% of those classified as hybrids should lack the line 
pattern. This feature is a secondary sexual character that 
may be more readily modified by hybridization than 
primary characters. 

The steep graduation of the tail occurs in both sexes 
of Sharp-tailed Grouse and is therefore not a secondary 
sexual character. The trends in measurements do not 
completely conform to the pattern shown in other char- 
acters. One might expect that in Class 5 (Table 4) in 
1964-1970 the influence of Prairie Sharp-tail genes 
might produce a more steeply graduated tail. Perhaps 
this unusual tail structure is not sufficiently integrated 
in the gene matrix of the species to resist the more con- 
ventional square tail of the Prairie Chicken genes in 
hybrids. 

Tables 2-4, 6 and 7 showed a number of trends. In 
the first period, 1949-1959, there was a tendency for 
hybrids to cluster toward Class | and 2 on the Prairie 
Chicken end of the scale. By the third period, 1964— 
1970, the clustering was toward Class 4 and 5 or the 
Sharp-tailed Grouse end of the scale. 

In Class 1 and 2 there was a strong tendency for the 
number of specimens to decline in each period while 
the number in Class 4 and 5 showed an increase. The 
latter is probably because Prairie Sharp-tailed Grouse 
had completely colonized the island and were con- 
tributing their genes to the mix. Observations indicated 
that the alpha males at Sheguiandah and at Billings lek 
in 1966 were probably Prairie Sharp-tailed Grouse 
(Lumsden 2005). 


LUMSDEN: PLUMAGE AND INTERNAL MORPHOLOGY OF ““PRAIRE GROUSE” 


523 


Ischiadic foramen 


Ischium 


Ficure 5. [lum process of (a) Greater Praire Chicken (b) and 
(c.) Heath Hen and (d) Northern Sharp-tailed Grouse. 


The speed with which change took place in the mor- 
phology of the Manitoulin population over time was 
perhaps largely due to the fact that “Prairie” Grouse are 
lek species. The particular bird that succeeds in becom- 
ing the alpha male in the hierarchy of dominance on a 
lek would consummate most of the matings. His genes 
would dominate in the subsequent year class. The 
absence of male Prairie Chickens on the Airport lek in 
1962 and the presence of 10 male Prairie Sharp-tailed 
Grouse would have had a profound effect on the 1962- 
1963 year class. It is perhaps not surprising that Guild 
(1997*) saw 50 Sharp-tailed Grouse there on 19 April 
1997. J. Iron (personal communication) saw 75-100 on 
15 April 2000, 30 on 12 April 2003 and 60 Sharp-tailed 
Grouse there on 11 April 2004. Those observers made 
no mention of Prairie Chickens or hybrids. Being skilled 
naturalists, they are unlikely to have missed them. 

Note: The extinct Heath Hen (7ympanuchus c. cupi- 
do) is of interest in that its ilium process does not proj- 
ect as far as that of the Greater Prairie Chicken (Tym- 


524 


panuchus c. pinnatus) (Figure 4). The two specimens 
I have examined would be placed in Class 2 and 3 of 
Table 7. Gutiérrez et al. (2000) judged the Heath Hen 
to be a species level taxon. The structure of the ilium 
may support that judgement. 


Acknowledgments 

I thank L. L Snyder, former Curator of Birds at the 
Royal Ontario Museum, for working space and encour- 
agement. For the loan of specimens I thank J. J. Hickey, 
Department of Wildlife Ecology, Madison Wisconsin; 
R. W. Storer, University of Michigan Museum, Ann 
Arbor; D. D Ripley, Peabody Museum, Yale; H. Fried- 
man, U.S. National Museum, Washington, D.C.; and 
the Field Museum, Chicago. 

Many individuals supplied information and speci- 
mens. My particular thanks go to W. Ritching, B. 
Smith, H. McQuarrie, H. Bailey and J. Budd. I thank 
F. and F. N. Hamerstrom, Wisconsin Department of 
Natural Resources, and A. Ammann, Michigan Depart- 
ment of National Resources, for valuable discussions 
and unpublished data. I received help collecting Prairie 
Chicken specimens from J. Horak, G. Smart, R. D. 
Buckingham, K. Mentzel, C. Baker, N. D. Patrick, L. 
Bluss, J. B. Dawson, B. Stephenson, R. Hepburn and 
many field staff from the Ontario Ministry of Natural 
Resources. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Documents Cited (marked * in text) 

Guild, J. 1997. Sharp-tailed Grouse at Gore Bay. Ontario 
Field Ornithologists News: page 7. 

Lumsden, H. G. 1965. Displays of the Sharp-tailed Grouse. 
Unpublished research report 66. Technical Series. Depart- 
ment of Lands and Forests: 68 pages. 


Literature Cited 

Campbell, B., and E. Lack. Editors. 1985. A dictionary of 
birds. 670 pages. 

Gutiérrez, J. R., G. F. Barrowclough, and J. G. Groth. 
2000. A classification of the grouse (Aves; Tetraoninae) : 
Based on mitochondrial DNA sequences. Wildlife Biology 
6: 205-211. 

Lumsden, H. G. 2005. “Prairie Grouse”: Hybridization on 
Manitoulin Island. Canadian Field-Naturalist 119: 507-514. 

Manweiler, J. 1939. The combined weight class-rectrix pat- 
tern method for determining sex of Sharp-tailed Grouse. 
Journal of Wildlife Management 3: 283-287. 

Schwartz, C. W. 1945. The ecology of the Prairie Chicken 
in Missouri. University of Missouri Studies 20. 99 pages. 

Snyder, L. L. 1935. A study of the Sharp-tailed Grouse: 
Contributions of the Royal Ontario Museum, University 
of Toronto Press: 66 pages. 


Received 15 April 2005 
Accepted 15 September 2005 


Ce! 


Response of Pale Swallow-wort, Vincetoxicum rossicum, following 
Aboveground Tissue Loss: Implications for the Timing of Mechanical 
Control 


Curtis I. MCKAGUE* and NAOMI CAPPUCCINO 


Department of Biology, Carleton University, Ottawa, Ontario K1S 5B6 Canada 
* Author for correspondence; Current address: Department of Integrative Biology, University of Guelph, Guelph, Ontario, 
NIG 2W1 Canada; Telephone: 519-824-4120 X 56307; e-mail address: cmckague @uoguelph.ca 


McKague, Curtis I., and Naomi Cappuccino. 2005. Response of Pale Swallow-wort, Vincetoxicum rossicum, following above- 
ground tissue loss: implications for the timing of mechanical control. Canadian Field-Naturalist 119(4): 525-531. 


The growth and reproduction of Pale Swallow-wort, Vincetoxicum rossicum, were investigated following differentially timed 
above-ground tissue loss by clipping throughout the growing season in Ottawa, Canada. If clipping occurred early in the 
growing season (before June), mature plants were able to compensate in height but not in biomass or reproductive output 
when compared to uncut controls. This compensation in height did not seem to come at the expense of below-ground storage 
tissue; there was no significant difference in root mass among the control and treatment conditions. Final plant height, mass, 
and reproduction declined as plants were cut later in the season. Pale Swallow-wort seedlings clipped before June were able 
to compensate in height when compared to uncut controls. Unlike mature plants, this compensation in height did come at the 
expense of root mass. As the clipping treatment was performed later into the growing season, both final plant height and 
shoot mass decreased, while root mass increased. The best time for controlling the population spread of Pale Swallow-wort 
using a single cutting treatment was on or near 26 June. Plants cut earlier than this date were able resprout and produce 
seeds; those cut after had already produced seeds that appeared viable and may be able to germinate if the cut stems were left 
in the field. When a single cutting or mowing treatment is to be employed for controlling Pale Swallow-wort, we recommend 
cutting after the first fruits are produced but before they are fully developed. 


Key Words: Pale Swallow-wort, Dog-strangling Vine, Vincetoxicum rossicum, Cynanchum rossicum, mechanical control, invasive 


plants, Ottawa. 


Pale Swallow-wort, Vincetoxicum rossicum Kleo. 
Barb. (Asclepiadaceae; synonym Cynanchum rossi- 
cum), also known as Dog-strangling Vine, is an intro- 
duced plant species that is becoming an increasingly 
problematic weed of natural areas in the northeastern 
United States and Ontario. Native to Ukraine and Rus- 
sia, Pale Swallow-wort was introduced into North 
America in the late 1800s (Sheeley and Raynal 1996). 
Pale Swallow-wort has been documented across south- 
ern Ontario from Toronto to Ottawa (DiTommaso et 
al. 2005), and has recently spread across the Ottawa 
River into the Outaouais region of Quebec and has been 
located in Montreal (DiTommaso et al. 2005). Within 
the United States, Pale Swallow-wort is widely distrib- 
uted within New York State, and has also been docu- 
mented in Connecticut, Indiana, Massachusetts, Michi- 
gan, New Hampshire, New Jersey, and Pennsylvania 
(DiTommaso et al. 2005). 

Pale Swallow-wort (Figure |) is a perennial herba- 
ceous vine, with stem lengths of mature individuals 
ranging from 60-250 cm (DiTommaso et al. 2005). 
Stems are often intertwined within dense patches while 
individual plants climb upon other vegetation for struc- 
tural support. Leaves are ovate and acute at the tip and 
are opposite in arrangement (DiTommaso et al. 2005). 
Flowers are pink to red in coloration, 5-7 mm in diam- 
eter, and have five fleshy petals arranged in a star con- 
figuration. Fruits, often arranged in pairs, are slender 


follicles 4-7 cm long (St. Denis and Cappuccino 2004; 
DiTommaso et al. 2005). In late summer, follicles re- 
lease wind-dispersed comose seeds which are 3-5 mm 
in length (DiTommaso et al. 2005). 

Pale Swallow-wort is capable of thriving under var- 
ious environmental conditions and has been document- 
ed in numerous habitat types, including lawns, gardens, 
old fields, disturbed areas, and along both coniferous 
and deciduous forest edges as well as within their 
understoreys (Sheeley and Raynal 1996; DiTommaso 
et al. 2005). Pale Swallow-wort thrives on limestone- 
based soils and is invading globally rare alvar habitats 
on the limestone plains of northern New York State 
(DiTommaso et al. 2005 and personal observation). 
Once established, populations of Pale Swallow-wort 
grow densely and spread aggressively, often eliminat- 
ing native plants (Sheeley and Raynal 1996; Chris- 
tensen 1998). Competitive displacement of native plant 
species by Pale Swallow-wort may have negative con- 
sequences for local faunal. Ernst and Cappuccino (2005) 
found that diversity and abundance of both stem- and 
ground-dwelling arthropod species were substantially 
lower in patches of Pale Swallow-wort than in patches 
of native old-field plant species (Asclepias syriaca, Sol- 
idago altissima, and mixed graminoids). Pale Swal- 
low-wort is also detrimental to the Monarch butterfly 
(Danaus plexippus) which has been observed oviposit- 
ing on it rather than on milkweed (Asclepias syriaca), 


325 


526 


its usual plant host, resulting in larval death (Mattila 
and Otis 2003). 

Although Pale Swallow-wort is not listed in any 
federal or provincial/state weed or seed acts in Canada 
or the United States (DiTommaso et al. 2005), it is 
recognized as a noxious weed and control programs are 
initiated at the municipal level. Attempting to control 
the spread of Pale Swallow-wort is a difficult task. It is 
self-compatible, so a single propagule can initiate a 
new infestation (St. Denis and Cappuccino 2004). 
Individual plants produce large numbers of viable 
seeds, which require neither dormancy nor stratifica- 
tion (Christensen 1998). Seedling survivorship is very 
high, ranging from 70 to 100% (Ladd and Cappuccino, 
in press). Seeds are polyembryonic (Sheeley and Ray- 
nal 1996; St. Denis and Cappuccino 2004), producing 
multiple seedlings about half the time (Cappuccino et 
al. 2002). 

Options for controlling Swallow-wort are limited. 
Herbicides such as glyphosate are moderately effective; 
however, several applications are needed (Christensen 
1998; Lawlor and Raynal 2002). A growing number of 
communities are banning herbicides due to their toxi- 
city to humans and non-target organisms (DiTomaso 
1997); in these places, manual means of controlling 
Swallow-wort are necessary. Repeated mowing can be 
an effective way to diminish the seed crop, but it does 
not kill the plants which resprout from buds at the root 
crown (Christensen 1998; DiTommaso et al. 2005). 
Moreover, mowing may not be feasible where equip- 
ment and labour are limiting. Mowing areas that are 
not monotypic stands of Pale Swallow-wort may alter 
the structure of the native plant community, which may 
be detrimental to insect populations (DiTomaso 1997). 

For smaller infestations of Pale Swallow-wort, 
selectively clipping individuals may be a more ecologi- 
cally effective solution. If available resources dictate 
that only a single cut is to be made, the timing of that 
cut is critical in determining whether seed production 
will be prevented. In general, plants clipped early in 
the growing season are more likely to compensate in 
growth and/or reproduction (Maschinski and Whitham 
1989; Bergelson and Crawley 1992; Mutikainen et al. 
1994; Bergelson et al. 1996; Lennartsson et al. 1998). 
If clipping occurs later in the growing season, plants 
often have insufficient time for compensatory growth. 
A late-season cut may thus seem preferable; however, 
seeds already produced by the plant may be viable if the 
cut stems are left at the site or scattered by a mower. 

The present study was performed to better under- 
stand the growth and reproductive responses of Pale 
Swallow-wort following aboveground tissue loss by 
clipping at different times throughout the growing sea- 
son. From this increased understanding in Pale Swal- 
low-wort biology, we hope that ecologically friendly 
and effective control methods may be used to slow the 
population growth of this species, thereby minimizing 
its deleterious effects on natural habitats. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Study Area and Methods 
Response of mature plants to differentially timed 
aboveground tissue loss 

This experiment was performed in an old field/for- 
est edge habitat in Hog’s Back Park, Ottawa, Ontario 
(45°22.45'N, 75°41.65'W), from May-August 2003. 
In mid-May, 150 individual plants of Vincetoxicum 
rossicum With initial heights of 30.2 + 0.2 cm (mean 
+ SE) were measured and randomly assigned to one 
of five differentially timed clipping treatments, or to 
the uncut control condition. Initially, 25 plants were 
assigned to each treatment; however, some plants were 
lost during the course of this experiment due to dam- 
age by humans or other animals. 

Beginning on 29 May, all plant heights were meas- 
ured and flower or fruit formation was noted. The first 
clipping treatment was performed this day. Clipping 
involved cutting the plants at ground level. Cut plants 
were dried for three days in a drying oven at 60°C and 
weighed. Subsequent clipping treatments were per- 
formed every two weeks until the final clipping treat- 
ment on 24 July. Plant heights were recorded on each 
clipping date and again on 7 and 21 August. Above- 
and below-ground tissues for all plants were harvested 
on 21 August and dried for three days in the drying 
oven. Roots and shoots were weighed separately for 
each plant. Any seeds that were produced were counted. 


Response of seedlings to differentially timed above- 
ground tissue loss 

A similar experiment was performed in an enclosed 
old field setting on Carleton University property in 
Ottawa, Ontario (45°23.023'N, 75°41.553"W) from 
May-August 2003. During April 2003, V. rossicum 
plants were grown from seed in a greenhouse at Car- 
leton University. In mid-May, 150 of these seedlings 
with initial heights of 18.0 + 0.4 cm (mean + SE) were 
transplanted into the old field after a two-week out- 
door acclimatization period. Seedlings were randomly 
assigned to the same clipping treatments described 
above for mature plants. 


Statistical Analyses 

All statistical analyses were performed using SPSS 
11.5 for Windows (standard base version). Pre-treat- 
ment heights for both adult plants and seedlings in 
the different treatments were compared using one-way 
ANOVAs. Because the assumptions of parametric sta- 
tistics were not met, Kruskal-Wallis tests were used to 
investigate among-treatment differences in final heights, 
shoot mass, and root mass for both adult plants and 
seedlings, as well as the seed output of adult plants. 
When a statistically significant difference was observed 
among treatments, a Games-Howell post-hoc test, 


which allows for unequal variance and unequal sample 


size, was performed to determine which treatments 
differed from one another (Games and Howell 1976). 


2005 


Results 
Response of mature plants to differentially timed 
above-ground tissue loss 

Pre-treatment heights did not differ significantly 
among the treatments (ANOVA, F 5,4, = 1.754, 
P = 0.126). The time of the growing season when 
plants were cut had a significant effect on final height 
(Kruskal-Wallis, H = 85.786 df = 5, P < 0.001). The 
Games-Howell post-hoc test revealed that plants that 
were cut on 29 May were able to compensate in height; 
they were not significantly different in height from the 
uncut controls (Figure 2). The final heights of plants 
cut on 29 May, 12 and 26 June were not significantly 
different from one another but they did differ from both 
10 and 24 July treatments. The 10 and 24 July treat- 
ment plants were not significantly different from one 
another but they did differ significantly from all other 
treatment groups in the experiment including uncut 
controls. 

The timing of the cut did not have a significant 
effect on root mass (Figure 3, Kruskal-Wallis, H = 6.123 
df = 5, P= 0.294), while shoot mass did differ signif- 
icantly among treatments (Kruskal-Wallis, H = 96.952 
df = 5, P < 0.001). The Games-Howell post-hoc test 
revealed that none of the experimentally clipped plant 
groups were able to compensate in shoot mass; all clip- 
ping treatments weighed significantly less than the un- 
cut control. Plants cut on 29 May, 12 and 26 June were 
not significantly different from one another but they 


McKAGUE AND CAPPUCCINO: RESPONSE OF PALE SWALLOW-WORT 


S27 


FIGURE |. Pale Swallow-wort, Vincetoxicum rossicum. A. 
An individual plant characteristically climbing sub- 
strate for structural support; B. Leaves and fruits; C. 
Inflorescences showing buds and mature flowers 
(Figure borrowed from DiTommaso et al. 2005, with 
permission from the authors and Canadian Journal 
of Plant Science). 


9 @) PRS ee eee ee es et ee et En Se ee ee ee ne re ee ee et te een tee ee ce 


80 
70 
60 


50 ab 


Height (cm) 


30 
20 
10 


0 


Control 29 May 


40 bp 


12 June 


10 July 24 July 


ao 


26 June 


Treatment Date 


FIGURE 2. Final height of mature Pale Swallow-wort plants clipped on various dates. Values are means + SE. Clipping date 
had a significant effect on plant mean height (Kruskal-Wallis test, H = 85.786, df = 5, P < 0.001). Ranked means 
that were significantly different (P < 0.05) according to Games and Howell’s (1976) multiple comparisons test are 


assigned different letters. 


Mass (g) 


—_ 


1S 
0.5 
@) si 


Control 


i 


29 May 


THE CANADIAN FIELD-NATURALIST 


12 June 
Treatment Date 


Vol. 119 


L] Root Mass 
Shoot Mass 


26 June 10 July 24 July 


FIGURE 3. Final root and shoot mass of mature Pale Swallow-wort plants clipped on various dates. Values are means + SE. 
Clipping date had a significant effect on mean shoot mass (Kruskal-Wallis test, H = 96.952, df = 5, P < 0.001) but 
there was no difference in mean root mass among treatments (Kruskal-Wallis test, H = 6.123, df = 5, P = 0.294), 
Ranked means that were significantly different (P < 0.05) according to Games and Howell’s (1976) multiple com- 


parisons test are assigned different letters. 


did differ from both 10 and 24 July treatments. The 10 
and 24 July treatment plants were not significantly 
different from one another but they did differ signifi- 
cantly from all other treatment groups. 

Seed production was greatly affected by clipping 
treatments (Table 1). Experimentally cut plants showed 
a significant decline in their reproductive output in com- 
parison to uncut controls (Kruskal-Wallis, H = 50.107 
df = 5, P < 0.001). The Games-Howell post-hoc test 
revealed that the clipping treatments were not signifi- 
cantly different from one another. No fruit produc- 
tion was observed until 26 June, and those plants that 
produced seeds despite having been clipped (29 May 
and 12 June cutting treatments) did so by 24 July and 
7 August, respectively. Plants cut on or after 26 June 
failed to produce any seeds. 


Response of seedlings to differentially timed above- 
ground tissue loss 

Seedlings in the six treatments did not differ sig- 
nificantly in initial height (ANOVA, F , ,4, = 0.480, 
P = 0.791). The timing of clipping had a significant 
effect on their final height (Kruskal-Wallis, H = 66.931 
df = 5, P < 0.001). The Games-Howell post-hoc test 
revealed that plants that were cut on 30 May were able 
to compensate in height as they were not significantly 
different in height from the uncut controls (Figure 4). 
Plants cut on 13 and 27 June, and 11 and 25 July were 
significantly smaller at the end of the season than those 
in the control treatment and the 30 May treatment. 

The time of the growing season when seedlings 
were cut had a significant effect on root mass (Figure 
5, Kruskal-Wallis, H = 21.013 df = 5, P = 0.001). 


TABLE |. Seed production of mature Pale Swallow-wort plants clipped on various dates. Number of seeds per plant are 
group means with SE in parentheses. Clipping date had a significant effect on reproductive output (Kruskal-Wallis test, H = 
50.107, df= 5, P < 0.001). Treatments with significantly different (P < 0.05) seed production according to Games and Howell's 
(1976) multiple comparisons test are assigned different letters. 


Proportion producing seed 


Treatment Group n 

Control P| 0.56 
29 May 24 O13 
12 June 22 0.14 
26 June 23 0.00 
10 July 24 0.00 
24 July 24 0.00 


Number of seeds per plant 


AT 2 (iS 2)P 
5,44 GSP 
2:3, (leaye 
Oe.) COLO}? 
0.0 (0.0)? 


COs OO? 


2005 McKAGUE AND CAPPUCCINO: RESPONSE OF PALE SWALLOW-WORT 529 


25 Be Pee ee Tee Reh SARE Ee MONE DMD. HORS. erme nT. 7! LAER ut Sutin tendons ddanemaden auton das doneucavavanavavantouuntonadancmvauadsenssvastoneiuvasnuctuvivecseneases 
a 
20 
a 
E 15 
& 
= 
a b 
2 10 
bc bc 
) Cc 
, a onl 


Control 30 May 13 June 27 June 11 July 25 July 


Treatment Date 


FiGuRE 4. Final height of Pale Swallow-wort seedlings clipped on various dates. Values are means + SE. Clipping date had 
a significant effect on plant mean height (Kruskal-Wallis test, H = 66.931, df =5, P < 0.001). Ranked means that 
were significantly different (P < 0.05) according to Games and Howell’s (1976) multiple comparisons test are 
assigned different letters. 


C1 Root Mass 


ge a Shoot Mass 


; ab ab 
Dog 
” 
” 
= 06 
0.4 
02 
Gc c & 
@) SN OR Se ee ee 
Control 30 May 13 June 2/7 June 11 July 25 July 
Treatment Date 


FIGURE 5. Final root and shoot mass of Pale Swallow-wort seedlings clipped on various dates. Values are means + SE. Clip- 
ping date had a significant effect on mean shoot mass (Kruskal-Wallis test, H = 68.644, df = 5, P < 0.001) and on 
mean root mass (Kruskal- Wallis test, H = 21.013, df = 5, P = 0.001). Ranked means of shoot mass that were signif- 
icantly different (P < 0.05) according to Games and Howell’s (1976) multiple comparisons test are assigned differ- 
ent uppercase letters while ranked means of root mass that were significantly different (P < 0.05) according to 
Games and Howell’s (1976) multiple comparisons test are assigned different lowercase letters. 


530 


Root mass was lowest for the 30 May treatment group 
and scores tended to increase in value as plants were 
cut later in the season. Final shoot mass also differed 
significantly among treatment groups (Figure 5, Krus- 
kal-Wallis, H = 68.644 df = 5, P < 0.001). The Games- 
Howell post-hoc test revealed that none of the experi- 
mentally clipped plant groups were able to compensate 
in shoot mass; all clipping treatments differed signifi- 
cantly from the controls. No seedlings produced flow- 
ers or seeds. 


Discussion 

When the stems of Pale Swallow-wort are cut dur- 
ing the growing season, new shoots emerge from buds 
at the root crown. In the present study, in which pri- 
mary shoots were cut at two-week intervals throughout 
the growing season, Pale Swallow-wort plants resprout- 
ed, but were not able to compensate fully for tissue 
loss. As we expected from the results of previous stud- 
ies on other plant species (e.g., Maschinski and Whit- 
ham 1989; Bergelson and Crawley 1992; Mutikainen 
et al. 1994; Bergelson et al. 1996; Lennartsson et al. 
1998), the ability of Pale Swallow-wort to partially 
compensate for tissue loss declined as plants were 
clipped progressively later in the growing season. AI- 
though plants cut on 29-30 May appeared to compen- 
sate in height, by the end of the growing season the 
new shoots had not attained the same biomass as the 
primary shoots produced by the uncut controls. 

While the root mass of mature Pale Swallow-wort 
plants showed no significant effect of above-ground tis- 
sue removal, the clipping treatment significantly dimin- 
ished the root mass of the seedlings. Seedlings lack the 
extensive root system of the mature plants, and their 
partial compensation in height and biomass seemed 
to come at the expense of their roots. All seedlings in 
our experiment survived until the end of August. How- 
ever, in a summer with less rainfall than 2003, the re- 
allocation of resources from root to shoot following 
clipping could leave the seedlings with underdeveloped 
root systems incapable of sustaining the plant through 
a drought (e.g., Bilbrough and Richards 1993; Perkins 
and Owens 2003). 

Mature control plants allocated nearly half their 
biomass to roots, a high proportion for mid- to late- 
successional species (Parrish and Bazzaz 1982). The 
high root:shoot ratio is partly explained by the weak- 
ness of the stem of this vine-forming plant; stems that 
have not twined around neighbouring vegetation gen- 
erally cannot support the weight of the fruits and flop 
to the ground once fruit production is underway. How- 


ever, the plant also possesses an extensive fibrous root 


system with a thick, woody rootstock, which likely con- 
tributes to its apparent ability to outcompete native 
vegetation and persist for decades (Sheeley and Ray- 
nal 1996). Seedling root:shoot ratios were even high- 
er; control seedlings invested over three times more tis- 
sue into root mass versus shoot mass. The seedlings 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


appeared to be developing below-ground storage tis- 
sue in preparation for future growing seasons. This may 
explain the slow maturation of Pale Swallow-wort; in 
a separate experiment, individuals planted as seeds four 
years ago have yet to flower (N. Cappuccino, unpub- 
lished data). 

Seed production was reduced dramatically by clip- 
ping. Clipping on or after 26 June halted seed pro- 
duction entirely. Clipping earlier in the season was 
insufficient to completely control reproductive output; 
over 10% of the plants clipped on 29 May and 12 June 
produced seeds. Fruits were first observed on 26 June 
in the controls and the three treatments that had not 
previously been cut. A single clipping treatment dur- 
ing this period of the growing season would be most 
effective because the fruits on this date contained only 
immature seeds that would be unable to germinate if 
left in the field following clipping. Plants clipped on 
or after 10 July also failed to produce seeds following 
clipping, but prior to clipping, their fruits contained 
seeds that appeared to be mature (they were filled and 
darkening in preparation for release). Although our 
data suggest that clipping in late June would be most 
effective, the timing of fruit production in other years 
and at other sites may vary. We recommend that plants 
be monitored for fruit formation and that cutting take 
place when the first fruits are formed but not fully 
developed. 

Mechanical control by clipping can be a success- 
ful method for controlling localized patches of Pale 
Swallow-wort that are threatening native species. How- 
ever, the ability of the plant to regrow following above- 
ground tissue loss both underscores the need for a more 
sustainable means of controlling this invasive weed, 
such as biological control, and argues against intro- 
ducing a leaf-chewing herbivore as the sole control 
agent. Adequate control of target plants following the 
introduction of a single herbivore is the exception 
rather than the rule (Julien 1989) and success is often 
achieved only after introduction of several herbivores 
(Denoth et al. 2002) that damage the plant in comple- 
mentary ways (James et al. 1992). Damaging the target 
plant in complementary ways can also be achieved 
through integrated weed management, for example, by 
combining mechanical control with biological control 
(Paynter and Flanagan 2004). Integrated weed manage- 
ment is increasingly being viewed as the most effec- 
tive strategy for controlling invasive plants in natural 
areas (Buckley et al. 2004) and is likely our best hope 
for controlling Pale Swallow-wort. 


Acknowledgments 

This research was funded by the Natural Sciences 
and Engineering Research Council of Canada (Under- 
graduate Student Research Award to CIM and Dis- 
covery Grant to NC). Permit for land access for this 
experiment was granted by Mike Muir of the National 
Capital Commission (NCC). The authors wish to thank 


2005 


the two anonymous reviewers for their helpful critical 
reviews of this manuscript; David Carpenter for his 
assistance in the field; Ed Bruggink for caring for our 
seedlings; and a special thanks to Jack and Yvonne 
McKague for their eternal support. 


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Received 6 July 2004 
Accepted 7 December 2005 


Differential Parental Care by Adult Mountain Plovers, Charadrius 
montanus 


STEPHEN J. DINSMORE!” and Fritz L. KNOPF? 


'Department of Wildlife and Fisheries, Mississippi State University, Box 9690, Mississippi State, Mississippi 39762 USA 

*Current address: Department of Natural Resource Ecology and Management, 339 Science II, Iowa State University, Ames, 
Iowa 50011 USA 

3U.S. Geological Survey, Fort Collins Science Center, 2150-C Centre Avenue, Fort Collins, Colorado 80526-8118 USA 


Dinsmore, Stephen J., and Fritz L. Knopf. 2005. Differential parental care by adult Mountain Plovers, Charadrius montanus. 
Canadian Field-Naturalist 1 19(4): 532-536. 


We studied chick survival of the Mountain Plover (Charadrius montanus) in Montana and found that chicks tended by females 
had higher survival rates than chicks tended by males, and that chick survival generally increased during the nesting season. 
Differences in chick survival were most pronounced early in the nesting season, and may be related to a larger sample of 
nests during this period. When compared to information about the nest survival of male- and female-tended plover nests, our 
chick data suggest a trade-off for adult plovers between the egg and chick phases of reproduction. Because Mountain Plover 
pairs have clutches at two nests at two different locations and show differential success between the sexes during the egg and 
chick phases, we offer that the Mountain Plover breeding system favours optimizing annual recruitment in a dynamic ecologic 
setting driven by annually unpredictable drought, grazing, and predation pressures. 


Key Words: Mountain Plover, Charadrius montanus, chick survival, Montana 


The Mountain Plover (Charadrius montanus) is an _ was related to the sex of the tending adult or varied sea- 
uncommon and locally distributed breeding bird of the — sonally. Here, we report estimates of daily chick sur- 
western Great Plains (Knopf 1996). The mating system, vival of Mountain Plovers in Montana and attempt to 
described as rapid multi-clutch (Graul 1973), is unusual —_ understand how survival is influenced by the sex of the 
in birds and involves two clutches per pair per year, tending adult and day within the nesting season. 
each clutch incubated by a single adult (Dinsmore 
2001). The female is thought to lay a complete clutch Methods 
for the male first, and then a second clutch at a differ- | Study area 
ent site for herself. The time between the start of incu- We studied Mountain Plovers in a 3000-km? area 
bation for each sex has not been well studied, but was in southern Phillips County in north-central Montana 
7-10 days for a small sample of nests in Montana (S.. (47°40'-47°55'N, 107°35'—108°30'W; Figure 1). The 
J. Dinsmore, personal observation). Because of this un- study area is bounded by the Missouri River to the 
usual mating system, Mountain Plovers offer an oppor- south, the Sun Prairie and Content roads to the east, 
tunity to investigate breeding strategy differences that Beaver Creek to the north, and Highway 191 to the west. 
may result from the sex of the tending adult. Earlier Approximately 2250 km? of the study area is in pub- 
work with this species has shown differential nest survi- lic ownership with the Bureau of Land Management 
val by sex of the incubating adult during the incubation (BLM, Malta Field Office) and the U. S. Fish and 
stage (Dinsmore et al. 2002); it is not known if these Wildlife Service (USFWS, Charles M. Russell National 
patterns extend through the chick phase to fledging. Wildlife Refuge). This area is a mixed-grass prairie with 

Although many aspects of Mountain Plover biology sagebrush (Artemisia spp.) flats bordering the south- 
have been well studied, there is little detailed infor- | western edge of the Prairie Pothole Region (Knowles 
mation about the survival of plover chicks. Miller and et al. 1982; Olson and Edge 1985; Dinsmore et al. 
Knopf (1993) estimated brood survival in northeastern 2002). All of our work took place on active Black-tailed 
Colorado and calculated a daily chick survival rate of | Prairie-dog (Cynomys ludovicianus) colonies that are 
0.979; the probability that a chick survived the 36- used preferentially by Mountain Plovers in Montana 
day fledgling period was 0.466. Knopf and Rupert (Knowles et al. 1982; Knowles and Knowles 1984; 
(1996) estimated the daily survival of plover chicks at | Dinsmore 2001). 
10-day age intervals and noted that survival generally 
increased with the age of the chick. Lukacs et al. (2004) 
estimated chick survival in east-central Colorado and © 
found that daily survival was lowest immediately after 
hatch, and quickly increased within 4 days post-hatch. 
None of those studies addressed whether chick survival 


Capture and marking 

During the 1999 nesting season, we fitted 28 nest- 
tending, adult Mountain Plovers with 3.0 g radio trans- 
mitters (Advanced Telemetry Systems, Isanti, Min- 
nesota) to allow us to monitor the survival of chicks 
within broods. Adult plovers were captured 1-4 days 


Nn 


a2 


2005 


prior to hatching date with a walk-in trap placed over 
the nest. On capture, we collected a feather sample from 
each adult for gender determination (see Dinsmore et 
al. 2003). Radio transmitters were glued to the mantle 
feathers of the adult using an epoxy (Titan Corporation, 
Lynnwood, Washington). Most adult plovers carried 
the transmitters until their chick(s) fledged and all trans- 
mitters were shed when the birds moulted prior to their 
fall migration. Chicks were marked with a unique series 
of four UV stable Darvic coloured leg bands (A. C. 
Hughes, London) and an aluminum U.S. Geological 
Survey band to allow individual identification. There 
were no adult or chick mortalities or injuries attribut- 
able to capture or handling. The Colorado State Uni- 
versity Animal Care and Use Committee approved the 
field methods used in this study (Protocol 98-134A-01). 

Using information from egg floatation (Dinsmore 
et al. 2002), we knew the exact hatch date for eggs in 
each nest and typically monitored chick survival at 1|- 
5 day intervals post-hatch. On each brood check, we 
located the adult bird using a hand-held Yagi antenna 
and observed the number of chicks and identity of each 
chick that was present. We ceased to monitor chicks 
when they fledged at an age of 33-36 days (see Knopf 
1996). 


Modeling chick survival 

We modeled the daily survival of plover chicks us- 
ing the nest survival model (Dinsmore et al. 2002) in 
Program MARK (White and Burnham 1999). This 
model is similar to the Kaplan-Meier model that is typ- 
ically used for telemetry data (Pollock et al. 1989), 
except that it does not require that the exact failure date 
be known. In our study, plover broods were not checked 
daily, so losses of individual chicks could only be as- 
signed to an interval. We used an estimate of 35 days 
as the fledging date, except in cases where we knew 
that the fledging date differed (but never by more than 
2 days). In a few cases, the tending adult lost its trans- 
mitter. In those cases, we censored the data at the last 
known brood check. 

In our analyses, we considered four models to ex- 
plain variation in chick survival of Mountain Plovers: 
(1) a model with constant survival [S(.)}, (2) a model 
where chick survival differed based on the sex of the 
tending adult [S (sex)], (3) model #2 plus an additive 
effect of a linear time trend [S (sex + T)], and (4) 
model #2 plus an additive effect of a quadratic time 
trend [S (sex + TT)]. The S(.) model is akin to a May- 
field estimate and implied that chick survival was 
constant across the entire nesting season, irrespective 
of date or sex of the tending adult. The S(sex) model 
implicitly allowed chick survival to differ based on the 
sex of the tending adult. We were uncertain whether 
this was an important influence on chick survival, but 
deemed it a relevant question based on significant dif- 
ferences in the incubation stage (Dinsmore et al. 2002). 
The S(sex + T) and S(sex + TT) models added sea- 
sonal variation in chick survival to the sex difference 


DINSMORE and KNOPF: PARENTAL CARE BY MOUNTAIN PLOVERS 


533 


Fort Belknap 
Incian Reservation 


hoe Cy ’ 
Willa fetes 


80 Kilometers 


FIGURE |. Map of Phillips County, Montana showing the dis- 
tribution in 2000 of Black-tailed Prairie Dog colonies. 
The stippled regions show an Indian Reservation and 
National Wildlife Reserve. The hatched portion is the 
study area. 


model. Based on our knowledge of plover biology, we 
suspected that chick survival might generally increase 
during the nesting season (Knopf and Rupert 1996), 
but were unsure whether this pattern might be linear 
or curvilinear. 

We assessed the fit of the nest survival model to our 
chick survival data using a chi-square test of independ- 
ence of the fates of chicks within broods as a function 
of brood size. We computed the expected values for the 
test statistic as the product of the number of broods in 
each clutch size (always two or three chicks) and the 
probability of observing that outcome. The latter prob- 
ability was calculated as the product of survival (# 
surviving/total) and mortality (# dying/total) within 
each possible brood outcome. For example, the prob- 
ability (Pr) of observing a brood of three chicks where 
only one survived 1s 3 * PatSneynin ey Pr(dying)*. We 
estimated over-dispersion as € = X~ and made this 
adjustment in MARK. oh 

We used the methodology of Burnham and Anderson 
(2002) to identify the best model(s) and make infer- 
ences about the factors influencing chick survival in 
plovers. We used Akaike’s Information Criterion (AIC; 
Akaike 1973), corrected for possible small sample bias 
and over-dispersion (QAICc), to rank the set of candi- 
date models. QAIC provides a means of objectively 
ranking a set of models and then selecting a “best ap- 


534 


proximating” model or models for inference (Burnham 
and Anderson 2002). We used QAICc values to com- 
pare the relative distances between the best approxi- 
mating model and each competing model. Generally, 
models with QAICc values <2 have strong support 
while those with QAICc values >10 have little support 
(Burnham and Anderson 2002). Normalized Akaike 
weights (w;) were also computed for each model to 
provide another measure of the relative importance 
of each model. Finally, we model averaged parameter 
estimates across all candidate models to obtain the 
“best” estimate(s) of chick survival (Burnham and 
Anderson 2002). 


Results 

We monitored 27 plover broods comprising 77 chicks 
(one adult plover died before its eggs hatched) during 
a 61-day period from 11 June to 10 August 1999. Two 
plovers (one male and one female) lost their transmit- 
ters in late June. Hatch dates varied, although we had 
slightly more broods early in the nesting season. The 
sample of tending adults was male-biased (18 males 
versus 9 females). 

The nest survival model fit our chick survival data 
well (2 = 7.08, P = 0.31), and our estimate of over- 
dispersion was ¢ = 1.18. We found good evidence for 
an effect of sex on brood survival (Table 1) with chicks 
in male-tended broods having lower daily survival 
(6. .= -0./8.0n a logit scale, SE = 0.42, 95% CI 
was —1.60, 0.05). This effect was always negative in 
the top three models. In addition to the sex effect on 
chick survival, we found evidence that survival varied 
during the nesting season (Table 1). The best model 
(w; = 0.53) had a quadratic time effect on survival, 
but a model with a linear effect on survival was near- 
ly as good (QAICc = 0. 29, w; = 0.46). Chick survival 
differed the most early in the nesting season, and after 
that was similar between male- and female-tended 
chicks. In both models, daily chick survival generally 
increased during the nesting season (Figures 2 and 3). 


Discussion 

Our results, while intriguing, should be interpreted 
with caution. By using information from all chicks 
within a brood, we risked violating the assumption of 
independent fates. It is important to note that violation 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


1 11 21 31 41 51 
Day 


FIGURE 2. Daily survival rates (95% CI) for Mountain Plover 
chicks tended by a male parent in Montana, 1999. 
Day | corresponds to 11 June. 


1 11 21 31 41 51 
Day 


FiGurE 3. Daily survival rates (95% CI) for Mountain Plover 
chicks tended by a female parent in Montana, 1999. 
Day | corresponds to 11 June. 


of the independence assumption does not affect the 
parameter estimates, but does provide estimates of pre- 
cision that are too small (Flint et al. 1995). Our test of 
independence of fates did not reveal any major viola- 
tion of this assumption, although our sample size of 
broods was small. However, our observations and pre- 
vious work with this species had shown that there was 
severe brood reduction within the first 1-2 weeks post- 
hatch (personal observation), but that many broods still 
fledged one or more chicks. Thus, attentiveness by the 
parent or some other factor was most likely the cause 
for chick losses, independent of the fates of other chicks 
in a brood. 

Each component of the avian life cycle must be fully 
understood before inferences can be made regarding 


TABLE |. Summary of model selection results for the chick survival of Mountain Plovers in southern Phillips County, Montana, 
1999. Models are ranked by ascending QAICc; w, is the model weight and K is the number of parameters. Daily chick sur- 
vival (S.) was modeled to include no time effects (.), a linear time trend (7), a quadratic time trend (77), and the effect of the 
gender of the tending adult (sex). QDeviance is computed as —2[log.(L(@)) - 2log.(L.(6))] where 9 represents a maximum 
likelihood estimate whose log-likelihood is evaluated for the model in question [L(@)] and for the saturated model [L.(0)]. 


Model QDeviance K 
S (sex + TT) 206.68 4 
S (sex + T) 208.99 3 
S (sex) 218.10 2 
SC) 234.16 l 


QAICe QAICc w, 
214.73 0.00 0.53 
215.02 0.29 0.46 
222.12 7.38 0.01 
236.16 21.43 0.00 


2005 


population processes. Information from some compo- 
nents (e.g., nest survival) is relatively easy to collect, 
but for others (e.g., chick and juvenile survival) it is 
more difficult, especially with precocial species like 
the Mountain Plover (see Lukacs et al. 2004). Collec- 
tively, such detailed information will present a clearer 
picture of the breeding cycle, and may hint at biolog- 
ical processes that influence success in each stage. 

In an earlier study (Dinsmore et al. 2002), we showed 
that male-tended Mountain Plover nests had much high- 
er survival during the incubation stage than did female- 
tended nests. We speculate that this results from some 
combination of temporal variation (male plovers are 
believed to tend the first nest), differential nest atten- 
tiveness, and the different physiological contributions 
of each sex (e.g., female plovers expend extra energy to 
produce eggs) early in the nesting cycle. Young Moun- 
tain Plovers are precocial (Knopf 1996) and typically 
leave the nest within hours of hatching, making it dif- 
ficult to ask questions about possible differences in the 
survival of chicks in male- and female-tended broods. 

Our results indicate that Mountain Plover chicks 
tended by the female have higher survival than those 
tended by the male, the opposite of what occurs during 
the incubation stage. Few birds share the same repro- 
ductive strategy as the Mountain Plover, and we were 
unable to find other published studies that addressed 
differential chick or brood survival as a result of the 
sex of the tending parent. A similar reproductive strat- 
egy has been well described for some species of North 
American quail (Burger et al. 1995; Pope and Craw- 
ford 2001), although these species differed from the 
Mountain Plover in adult sex ratio, nest initiation dates, 
and mean clutch size. 

So what does this mean? Male-tended Mountain 
Plover nests are more successful than female-tended 
nests, which result in a greater number of broods with 
a male parent. But during the fledgling stage, female- 
tended chicks have greater survival, even though there 
are fewer female-tended broods (Dinsmore et al. 2002). 
The ultimate cause(s) for this apparent trade-off are 
unknown, although we can pose two possible expla- 
nations. For successful early nests, which tend to be 
male-tended (Dinsmore et al. 2002), the adult male 
may still be engaged in courtship activities with later 
nesting females. This could result in a drop in brood 
attentiveness, and a resulting decline in the survival 
of chicks in male-tended broods. The reverse pattern 
would not be expected for females because a female 
with a brood may lack the physiological capability to 
produce an additional clutch. 

A second explanation, not necessarily independent 
of the first, attempts to reconcile the parenting conflicts 
of male plovers. Despite the asynchrony in nest initi- 
ation between the sexes, it appears that both members 
of the pair forage together during the incubation period 
(Graul 1973). In this scenario, the male may be in a 
conflict situation as he enters brood rearing while the 
female is still incubating. His attempts to maintain the 


DINSMORE and KNOPF: PARENTAL CARE BY MOUNTAIN PLOVERS 


333 


pair bond might initially compromise his attentiveness 
to newly hatched chicks, resulting in lowered chick 
survival early in the brood stage. Larger differences in 
chick survival early in the nesting season may be a result 
of our larger sample of broods during this, the peak 
period of brood rearing. 

The major evolutionary question that is still unan- 
swered is, “What are the relative contributions of each 
sex to annual recruitment?” On the basis of this and 
other studies of the Mountain Plover, we conclude that 
male and female plovers attain similar rates of produc- 
tivity with male plovers having higher nest survival and 
lower chick survival and female plovers showing a 
reverse pattern. These opposite patterns likely favor 
sustained productivity in the climatically unpredictable 
drought cycles of the Northern Great Plains. Putting the 
clutch of six eggs in two different nests at two different 
sites, each raised by a different adult, speaks to the flexi- 
bility in this breeding system that may have evolved in 
a highly unpredictable ecological landscape subject 
to major drought cycles, intensive grazing, and fluc- 
tuating predator populations. 


Acknowledgments 

We thank Fort Carson Military Reservation (U. S. 
Department of Defense), the U. S. Fish and Wildlife 
Service (Colorado Fish and Wildlife Management 
Assistance Office), the Biological Resources Divi- 
sion of the U. S. Geological Survey, the U. S. Bureau 
of Land Management (Malta Field Office, Montana), 
and the Denver Zoological Foundation for financial 
support. D. C. Ely, J. J. Grensten, and T. Hanks pro- 
vided field assistance. G. C. White provided statistical 
advice. The manuscript benefited from careful reviews 
by A. J. Erskine and J. P. Goossen. This is contribution 
number WF207 of the Forest and Wildlife Research 
Center, Mississippi State University. 


Literature Cited 

Akaike, H. 1973. Pages 267-281 in Information theory and 
an extension of the maximum likelihood principle. Edited 
by B. N. Petran and F. Csaki. International symposium on 
information theory. Second edition. Budapest, Hungary. 

Burger, L. W., Jr., M. R. Ryan, T. V. Dailey, and E. W. 
Kurzejeski. 1995. Reproductive strategies, success, and 
mating systems of Northern Bobwhite in Missouri. Jour- 
nal of Wildlife Management 59: 417-426. 

Burnham, K. P., and D. R. Anderson. 2002. Model selection 
and multimodel inference: a practical information-theoretic 
approach. Springer-Verlag, New York, New York. 

Dinsmore, S. J. 2001. Population biology of Mountain Plovers 
in southern Phillips County, Montana. Ph.D. dissertation, 
Colorado State University. 109 pages. 

Dinsmore, S. J., G. C. White, and F. L. Knopf. 2002. Ad- 
vanced techniques for modeling avian nest survival. Ecol- 
ogy 83: 3476-3488. 

Dinsmore, S. J., G. C. White, and F. L. Knopf. 2003. Annual 
survival and population estimates of Mountain Plovers in 
southern Phillips County, Montana. Ecological Applications 
13: 1013-1026. 


536 


Flint, P. L., K. H. Pollock, D. Thomas, and J. R. Sedinger. 
1995. Estimating prefledging survival: Allowing for brood 
mixing and dependence among brood mates. Journal of 
Wildlife Management 59: 448-455. 

Graul, W. D. 1973. Adaptive aspects of the Mountain Plover 
social system. The Living Bird 12: 69-94. 

Knopf, F. L. 1996. Mountain Plover (Charadrius mon- 
tanus). In The Birds of North America, Number 211. 
Edited by A. Poole and F. Gill. The Academy of Natural 
Sciences, Philadelphia, Pennsylvania, and The American 
Ornithologists’ Union, Washington D. C. 

Knopf, F. L., and J. R. Rupert. 1996. Reproduction and 
movements of Mountain Plovers breeding in Colorado. 
Wilson Bulletin 108: 28-35. 

Knowles, C. J., C. J. Stoner, and S. P. Gieb. 1982. Selective 
use of black-tailed prairie dog towns by Mountain Plovers. 
Condor 84: 71-74. 

Knowles, C. J., and P. R. Knowles. 1984. Additional records 
of Mountain Plovers using prairie dog towns in Montana. 
Prairie Naturalist 16: 183-186. 

Lukacs, P. M., V. J. Dreitz, F. L. Knopf, and K. P. Burn- 
ham. 2004. Estimating survival probabilities of unmarked 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


dependent young when detection is imperfect. Condor 
106: 927-932. 

Miller, B. J., and F. L. Knopf. 1993. Growth and survival 
of Mountain Plovers. Journal of Field Ornithology 64: 500- 
506. 

Olson, S. L., and D. Edge. 1985. Nest site selection by Moun- 
tain Plovers in Northcentral Montana. Journal of Range 
Management 38: 280-282. 

Pollock, K. H., S. R. Winterstein, C. M. Bunck, and D. D. 
Curtis. 1989. Survival analysis in telemetry studies: the 
staggered entry design. Journal of Wildlife Management 
Soe ay 

Pope, M. D., and J. A. Crawford. 2001. Male incubation and 
biparental care in Mountain Quail. Condor 103: 865-870. 

White, G. C., and K. P. Burnham. 1999. Program MARK: 
survival estimation from populations of marked animals. 
Bird Study 46: 120-139. 


Received 1 November 2004 
Accepted 23 September 2005 


Body Size Distribution and Frequency of Anthropogenic Injuries 
of Bluntnose Sixgill Sharks, Hexanchus griseus, at Flora Islets, 
British Columbia 


ROBERT DUNBRACK!*? and ROBERT ZIELINSKI 


'Biology Department, Memorial University, St. John, Newfoundland and Labrador A1B 3X9 Canada 
"Hornby Island Diving, Ford Cove, British Columbia VOR 1Z0 Canada 
3Corresponding author, e-mail: dunbrack@ mun.ca. 


Dunbrack, Robert, and Robert Zielinski. 2005. Body size distribution and frequency of anthropogenic injuries of Bluntnose 
Sixgill Sharks, Hexanchus griseus, at Flora Islets, British Columbia. The Canadian Field-Naturalist 119(4): 537-540. 


The Bluntnose Sixgill Shark (Hexanchus griseus) is a widely distributed demersal species whose population biology is poorly 
understood. Although H. griseus is normally found in deep continental slope waters, individuals from a population in the 
Strait of Georgia, British Columbia, make unexpected diurnal movements onto a shallow reef (Flora Islets) between June and 
August. This shallow water activity allowed in situ length measurements to be made on 35 free-swimming Bluntnose Sixgill 
Sharks using stereo videography. The measured sharks were all large juveniles and sub-adults, although smaller juveniles 
and pregnant females are known to occur in deeper adjacent waters. The restricted size distribution at Flora Islets may arise 
because small juveniles avoid contact with larger conspecifics and mating takes place offshore. All measured sharks were 
individually identified by unique scar patterns. In 13 of 35 sharks these scars were consistent with injuries expected from 


hooking and entanglement by commercial fishing gear. 


Key Words: Sixgill Sharks, Hexanchus griseus, length-frequency distribution, anthropogenic injury, British Columbia. 


The Bluntnose Sixgill Shark (Hexanchus griseus) 
is a demersal species found along the continental slopes 
of the Pacific, Atlantic, and Indian Oceans (Compagno 
1984). Hexanchus griseus is one of the most widely 
distributed (Compagno 1984) and largest of fishes 
(length to at least 4.8 m; Castro 1983), and the highest 
trophic level predator throughout its range (Ebert 1994; 
Froese and Pauly 2005"), feeding on a variety of large 
prey items (Hart 1973; Compagno 1984; Ebert 1986, 
1994). In British Columbia the Bluntnose Sixgill Shark 
occurs in the Strait of Georgia and the deep inlets of 
the mainland coast and the west coast of Vancouver 
Island (Hart 1973). There is little information on the 
population biology of H. griseus, despite its wide dis- 
tribution, because it is not usually exploited commer- 
cially and scientific collecting has been limited due to 
the potential for adverse effects on local populations 
(Ebert 1986, 1994; Clark and Kristof 1990; Carey and 
Clark 1995). Although H. griseus is normally a deep 
water species, in the vicinity of Flora Islets in the Strait 
of Georgia, British Columbia, a number of Bluntnose 
Sixgill Sharks make diurnal movements into shallow 
water (20-40 m) between June and August. This activ- 
ity provides a unique opportunity to collect quantitative 
behavioural and population data for H. griseus using 
simple and non-invasive underwater camera systems 
(Dunbrack and Zielinski 2003). In this paper a stereo 
video measurement technique is used to derive the body 
length-frequency relationship for Bluntnose Sixgill 
Sharks at Flora Islets. 


Study Area 

The study site is immediately adjacent to Flora Is- 
lets, a chain of three small islets southeast of Hornby 
Island in the Strait of Georgia (49°30.9'N, 124°34.5'W; 
Dunbrack and Zielinski 2003). The remote cameras 
used for length measurements were attached to the ver- 
tical face of a submerged rocky reef which parallels 
Flora Islets to the southwest 50 to 100 m offshore. 
Water depth at the base of the wall increases in a south- 
easterly direction from 30 m to over 200 m. 


Methods 

Length measurements were derived from analysis 
of paired images taken with two low light, black and 
white video cameras in waterproof housings. The cam- 
eras were mounted along a vertical pipe tied into the 
face of the reef wall with one camera 2.5 m above the 
other and both cameras facing down to the base of the 
wall which was 5 m below the lower camera and at a 
depth of 35 m. Individual cables from each camera ran 
to a surface float and terminated in waterproof con- 
nectors that could be plugged into leads on a surface 
vessel for recording. Stereo video length measurements 
of a 213 cm pipe, replicated in various parts of the cam- 
eras’ visual fields, had a mean error of 2% (maximum 
error 4.9%). A detailed description of the stereo meas- 
urement system is given in Dunbrack (in press). 

Individual sharks could be identified by distinctive 
lateral scar patterns; however, as a shark swam below 
the cameras, only its right or left side was visible. To 


537 


538 


ensure that individual sharks were included only once 
in the analysis, length measurements were restricted 
to sharks displaying their right side scar patterns, that 
is sharks moving from left to right, inbound from deep 
water (a more frequently observed pattern than right to 
left, outbound movements). All sharks that were com- 
pletely visible from both cameras, and could be indi- 
vidually identified, were measured. There was no body 
size bias in the ability to identify individuals. This 
was primarily dependent on the ambient light level, 
which was generally low within the two hours follow- 
ing sunrise or preceding sunset, and varied substan- 
tially in response to fluctuations in the density of par- 
ticulate matter in the upper water layers. 


Results 

A total of 50 length measurements of 35 individual 
sharks were obtained from recordings made between 
10 August and 15 September 2001 and between 10 July 
and 21 July 2002. The total of 50 includes one or two 
repeat measurements of 13 individuals that were filmed 
on multiple occasions. The average difference for these 
repeat measurements ((largest-smallest)/smallest) x 100) 
was 1.9% (maximum difference 4.7%). The body 
length-frequency distribution is roughly bell shaped 
with 80% of measurements falling between 180 cm 
and 300 cm (Figure 1). Mean, minimum and maximum 
lengths of the 35 sharks were 240 cm, 135 cm, and 
353 cm, respectively. In most cases it was not possible 
to determine a shark’s sex; however, all individuals 
greater than 280 cm were observed at close range 
under good illumination and could be sexed based on 
pelvic fin morphology. All sharks over 280 cm were 
females (n = 7). 

The scars used to identify sharks varied from rela- 
tively small light patches to groups of long parallel 


Figure 1 


Number 


Qo 1°72) 522] © L*2) [°2) 1°72) o 
wo ™ Qo oO © cor) N wo 
= it o nN N N ° oo 
v oO oO © oO oOo oO oO 
wo co ee: wz ™ oO oO 
= pee N N N Oo oO 

Length (cm) 


FIGURE |. Body length-frequency relationship for 35 Blunt- 
nose Sixgill Sharks observed at the Flora Islets reef. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


lines running along the right lateral dorsal surface 
(Figure 2). The origin of these parallel scars is uncer- 
tain but they are consistent with injuries expected from 
hooking and subsequent entanglement with commer- 
cial fishing gear (dogfish longlines or bait lines attached 
to commercial prawn traps). Scars of this type were 
seen in 13 of the 35 sharks, a number that probably 
underestimates injury frequency because observations 
were confined to the right hand dorsal side; the left side 
and the jaw region, where divers frequently observe 
hooking injuries, were not visible in any of the meas- 
ured sharks. 


Discussion 

Newborn Bluntnose Sixgill Sharks are approxi- 
mately 70 cm in length; minimum length at maturity 
is greater than 400 cm for females and greater than 
314 cm for males (Ebert 1989). The size range at Flora 
Islets was 135 cm to 353 cm and all sharks over 280 cm 
were females, indicating that at Flora Islets all Blunt- 
nose Sixgill Sharks are large juveniles and subadults. 
Smaller juveniles (75 cm) occur in deeper adjacent 
waters (Miller and Greenfield 1965) and there is a 
single record of a pregnant female (>400 cm) from 
the same area (S. McFarlane, Canada Fisheries and 
Oceans, Nanaimo, British Columbia, personal com- 
munication) The absence of adults and the smallest 
juveniles from Flora Islets reef is in agreement with 
data from Barkley Sound on the west coast of Van- 
couver Island. Here, visual length estimates of 55 H. 
griseus observed in shallow water by divers, ranged 
from 100 cm to a maximum of 300 cm for males and 
360 cm for females (N. McDaniel, Subsea Enterprises, 
Vancouver, British Columbia, personal communica- 
tion), whereas 43 immature sharks captured in the same 
area on deepwater longlines during a 1994 tagging 
study, varied in length from 86 cm to 400 cm (Canada 
Fisheries and Oceans, Nanaimo, British Columbia, 
unpublished report). These data indicate that in Barkley 
Sound the smallest juveniles probably stay in deep 
water, and that sexually mature fish are rare. 

The apparent absence of the smallest juveniles from 
shallow water in Barkley Sound and at Flora Islets 
could result from their use of habitats or foraging 
behaviors different from those of larger conspecifics, 
possibly to enhance growth or deter cannibalism (Ebert 
1989). The virtual lack of sexually mature adults in 
deep or shallow water at either site is problematic but 
could be connected to age-dependent movements 
whereby birth and juvenile development take place in 
productive inshore waters but mating occurs elsewhere, 
probably offshore. Geographic separation of adult and 
juvenile H. griseus has been reported previously (Ebert 
1989: Castro et al. 1999) and is common in other shark 
species (Hoenig and Gruber 1990; Compagno 2001). 

Approximately one third of the Bluntnose Sixgill 
Sharks observed at Flora Islets bore scars consistent 
with commercial gear entanglement; however, no in- 


2005 


juries of this type were observed in the seven sharks 
greater than 280 cm or in the three smallest sharks 
(<180 cm). Although the sample size is small, this 
suggests that the largest sharks may be able to break 
off hooks or leaders without entanglement, whereas 
sharks below 280 cm may escape after a period of 
struggle, but with a probability that decreases as body 
size decreases. Unfortunately, there are no bycatch 
data available for H. griseus to test this hypothesis. 

Body size-frequency distributions can provide infor- 
mation on a range of population-level processes in 
fishes including growth and mortality rates and size 
or age-based differences in habitat use (Moyle and 
Cech 1996). Length-frequency data for commercial 
species are readily available from catch statistics, and 
for non-commercial species are generally obtained 
from targeted collecting. However for large, deep water 
species, such as H. griseus, the collection of length- 
frequency data can be difficult or unacceptably destruc- 
tive and the size structure of most large deep water 
fishes is poorly known (Casey and Taniuchi 1990). 
Here we have demonstrated that meaningful popula- 
tion data for such species can be obtained using non- 
invasive remote imaging techniques applied at strate- 
gically placed observing stations. Although extension 
of this technique to other species would require the 
identification of similar seasonal population aggrega- 
tions, such seasonal movements are common in fishes. 
For example, we have recently used remote video imag- 
ing to quantify seasonal activity patterns in another 
little known deep water cartilaginous fish, the chimaerid 
Hydrolagus colliei (unpublished). 


Acknowledgments 

A. Heath, L. Giguere, N. McDaniel, B. Parsons, 
and RD’s family members provided valuable assistance 
at different points in this study. Two anonymous re- 
viewers made helpful comments on an earlier draft of 
the manuscript. This work was carried out with the co- 
operation of the Province of British Columbia Parks 
Branch, and was supported by Memorial University 
of Newfoundland and NSERC Canada grants to R. 
Dunbrack and M. Ramsay. 


Documents Cited (marked * in text) 

Froese, R., and D. Pauly. Editors. 2005. FishBase. World 
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Carey, F. G., and E. Clark. 1995. Depth telemetry from the 
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Casey, J. G., and T. Taniuchi. 1990. Shark tagging. Pages 
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National Oceanic and Atmospheric Administration Tech- 
nical Report National Marine Fisheries Service 90. 


DUNBRACK AND ZIELINSKI: BLUNTNOSE SIXGILL SHARKS 


339 


FIGURE 2. Hexanchus griseus outlines traced from single 
video frames. Injury markings are indicated by dark 
lines (injuries actually appear as lighter markings 
against a uniformly dark dorsal surface.) The top 
four sharks were viewed from their right side and 
show injuries along their lateral dorsal surface. The 
bottom shark (not measured) has a large indentation 
in its lower jaw caused by a hook. 


Castro, J. I. 1983. The sharks of North American waters. 
Texas A&M University Press, College Station, Texas, 
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Castro, J. I., C. M. Woodley, and R. L. Brudek. 1999. A 
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Clark, E., and E. Kristof. 1990. Deep-sea elasmobranchs 
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Received 2 November 2004 
Accepted 7 October 2005 


Use of Radio-Telemetry to Test for Investigator Effects on Nesting 
Mallards, Anas platyrhynchos 


TERRI D. THORN!, ROBERT B. EMERY”, DAVID W. HOWERTER, JAMES H. DEVRIES, and BRIAN L. JOYNT 


Institute for Wetland and Waterfowl Research, Ducks Unlimited Canada, P.O. Box 1160, Stonewall, Manitoba ROC 2Z0 
Canada 

‘Current Address: U.S. Fish and Wildlife Service, 3425 Miriam Ave., Bismarck, North Dakota 58501-7926 USA 

*Corresponding author: e-mail: b_emery @ducks.ca 


Thorn, Terri D., Robert B. Emery, David W. Howerter, James H. Devries, and Brian L. Joynt. 2005. Use of radio-telemetry to 
test for investigator effects on nesting Mallards, Anas platyrhynchos. Canadian Field-Naturalist 119(4): 541-545. 


We examined the effects of investigator activity on hatching rates of radio-marked wild female Mallards (Anas platyrhynchos), 
and evaluated the efficacy of radio-telemetry to minimize nest disturbance, characterize vegetation at nest sites, and mark nests 
for later relocation. Differences in hatching rates between birds that were flushed once (experimental) and those never flushed 
(control) approached significance (P = 0.086). However, hatching rates did not differ (P = 0.588) between the two groups 
when nests where investigator activity caused abandonment (30% of experimental nests) were removed from analysis. If the nest 
remained active, flushing the bird and visiting the nest once did not appear to increase the likelihood of the nest being depre- 
dated. We were able to locate 92% of radio-telemetry marked nests (control) after the female was no longer tending the nest. 
Radio-telemetry provides a technique to collect relatively unbiased nesting data for Mallards without disturbing the female. 


Key Words: Mallard, Anas platyrhynchos, nests, hatching-rates, investigator effects, abandonment, radio-telemetry. 


Investigating reproductive performance of waterfowl 
often requires physically locating nests followed by 
repeat visits until the nest’s fate is determined. Howev- 
er, locating and revisiting nesting birds may influence 
reproductive performance by causing nest abandonment 
and by subsequently affecting renesting. Also, inves- 
tigator visits to the nest may aid predators in locating 
and destroying nests and thus result in biased estimates 
of nest survival. Alternatively, predators may avoid 
human scent, leading to increased nest survival. 

Potential negative influences of investigator activity 
on estimates of reproductive success have been recog- 
nized for decades in waterfowl research (Bennett 1938; 
Low 1940; Hammond and Forward 1956). Unfortu- 
nately, results of studies of investigator impact on water- 
fowl hatching rates have been equivocal. Earl (1950) 
and Hammond and Forward (1956), suggested that 
investigator presence lowered nesting success, where- 
as Gotmark and Ahlund (1984), Livezey (1980), and 
Sedinger (1990) did not find this to be true. In contrast, 
observer activity may increase nest abandonment (Balat 
1969; Reed 1975; Livezey 1980). 

Various methods (see Gotmark 1992 for review) 
have been used to test for investigator influences on 
reproductive parameters. However, locating and mon- 
itoring same-aged, undisturbed, natural duck nests (con- 
trol) has been difficult. This can be crucial when trying 
to estimate accurately investigator-caused influences on 
hatching rates. Artificial nests have been used to facil- 
itate such comparisons (e.g., Gottfried and Thompson 
1978; Vacca and Handel 1988; Sullivan and Dinsmore 
1990). Extrapolating estimates from artificial nests to 


natural nests might not be entirely valid, because arti- 
ficial nests are “visited” when deployed (Gotmark et al. 
1990; MaclIvor et al. 1990; Hendricks and Reinking 
1994; but see Olson and Rohwer 1998), and artificial 
nests may be poor substitutes for estimating hatching 
rates of natural nests (see Butler and Rotella 1998). 
Radio-telemetry offers a partial alternative by allow- 
ing remote monitoring of nesting waterfowl. 

We used radio-telemetry to monitor duck reproduc- 
tive performance in response to upland habitat manip- 
ulation during the Prairie Habitat Joint Venture (PHJV) 
assessment program of the North American Waterfowl 
Management Plan. Original PHJV assessment program 
protocols stated that if a female was suspected of hav- 
ing a nest, she was flushed to determine nest site loca- 
tion. We designed a study to determine whether flushing 
females and visiting a nest once reduced hatching rates. 
Specifically, our objectives were to: (1) determine the 
impact on hatching rates of flushing a female Mallard 
(Anas platyrhynchos) and visiting her nest early in the 
nesting cycle, (2) if there was an impact, determine its 
cause, and (3) evaluate the efficacy of our radio-teleme- 
try nest-marking protocols. 


Study Areas and Methods 

Data were collected from three 64.75-km/? areas in 
the prairie pothole region of Canada (Bellrose 1980; 
Poston et al. 1990). Study areas were centred approx- 
imately 1 km west of Belmont, Manitoba (49°25'N, 
99°29'W), 5 km northeast of Davis, Saskatchewan 
(53°9'N, 105°37'W), and approximately 3 km west of 
Erskine, Alberta (52°19'N, 112°55'W). These landscapes, 


541 


542 


characterized by flat to rolling topography, have been 
altered by cultivation for cereal and oil-seed crops, 
forage production, and livestock grazing. 

We decoy-trapped (Sharp and Lokemoen 1987; 
Ringelman 1990) 135, 136 and 135 pre-laying female 
Mallards at Belmont, Davis and Erskine, respectively, 
during 6 — 28 April 1994. Females either were implant- 
ed with a radio transmitter (Olsen et al. 1992; all fe- 
males at Belmont and Davis and 67 females at Erskine) 
or transmitters were anchor/sutured to their backs 
(Mauser and Jarvis 1991; Pietz et al. 1995; 68 females 
at Erskine). After marking, we located birds twice 
daily between 06:00 and 13:00 (Gloutney et al. 1993) 
using vehicle-mounted, null-array antenna systems and 
triangulation. Nesting attempts were suspected when 
triangulation placed a female in the same location for 
5 consecutive mornings. We randomly allocated a sub- 
sample of all nesting attempts we encountered between 
6 May and 19 June to either a control or an experi- 
mental group. 

In the control group, nests were located and moni- 
tored only via telemetry; females were not flushed and 
nests were not visited while active. To estimate the 
location of control group nests we used hand-held 
antennas to take multiple bearings (< 5) around the 
probable nest site. The number of bearings and the 
distance away from the nest from which these bear- 
ings were taken varied with the complexity of the 
habitat type. For example, in very simple habitat con- 
figurations such as a small, isolated patch of nesting 
cover surrounded by sparse vegetation not suitable for 
nesting, we remained quite far (e.g. 100 — 150 m) from 
the nest and took only a few bearings. We aligned bear- 
ings with natural land features and/or small pieces of 
vinyl! flagging and drew a detailed map of the estimat- 
ed nest location to assist in relocating the nest when it 
was no longer active. Nesting site characteristics (habi- 
tat and vegetation type) were assigned to each nest 
from a distance. After a female Mallard was believed 
to be nesting, she was located via telemetry at least once 
each morning to monitor the status of the nest. If she 
was absent from the nest, a second location was ob- 
tained later the same day. If two consecutive teleme- 
try readings indicated that the female was not in the 
area of the nest, we immediately searched for the nest, 
recorded nest-site vegetation characteristics, and deter- 
mined the nest’s fate (Klett et al. 1986). 

In the experimental group, we visited the suspect- 
ed nest site location and flushed the bird after trian- 
gulation placed her in the same location for five con- 
secutive days. For each nest, we recorded vegetation 
characteristics around the nest site, number of eggs, 
and incubation stage (Weller 1956). We assumed a 
laying interval of one egg per day. Nest status was then 
monitored remotely using telemetry, and only when 
the female was absent for two consecutive telemetry 
locations was the nest visited again to determine its 
fate. Nest fate was considered abandoned due to inves- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


tigator activity when the stage of nesting (number of 
eggs or stage of incubation) did not change between 
nest visits (Klett et al. 1986) and subsequent telemetry 
locations indicated that the female was still alive but 
had failed to return to the nest after we had flushed her. 

Because the study was a part of the PHJV assess- 
ment program, where one of the main objectives was 
to determine waterfowl] nesting habitat preferences, we 
risked misclassifying control-group females as nesting 
when they were not. To minimize this risk to our over- 
all study objectives, we excluded from both groups 
those nesting attempts where nests were located < 30 m 
from wetlands with standing water. 

Vegetation characteristics around the nest site were 
recorded at two different scales. The habitat within a 
2-m radius around the nest was recorded as: (1) grass- 
land, (2) hayland, (3) planted cover, (4) cropland, (5) 
woodland, (6) shrubland, or (7) wetland (Emery et al. 
2005). These classifications were applied to any 
patch of habitat 210 m across in its narrowest dimen- 
sion. Nest site vegetation within a 0.5-m radius of the 
nest bowl was characterized as one or more of the 
following types: (0) unvegetated, (1) annual crop, (2) 
upland graminoids-native, (3) upland graminoids- 
introduced, (4) upland forbs, (5) low shrubs (<1 m), 
(6) tall shrubs (1 — 6 m), (7) trees, (8) short emergent 
hydrophytes (< 0.5 m), and (9) medium/tall hydrophytes 
(>0.5 m). 


Data Analysis 

To provide adequate sample sizes for analyses, data 
were pooled across habitats, study areas, and trans- 
mitter types. A chi-square test (PROC FREQ; SAS 
Institute 1997) was used to determine if there was a 
difference in the probability of hatching at least one 
egg (successful nest) between females that were flushed 
and those that were not. To test for investigator effects 
other than nest abandonment (i.e., predation), we re- 
moved those nests in the experimental group where 
investigators caused abandonment (Klett et al. 1986) 
and used Johnson’s (1979) modification to the May- 
field method (Mayfield 1961, 1975) to estimate daily 
survival rates and hatching rates for the two groups. To 
detect differences in hatching rates, daily survival rates, 
weighted by exposure days, were used in a least-squares 
linear model (PROC GLM; SAS Institute 1997; Green- 
wood et al. 1995). To test for potential negative effects 
of radio-transmitters we compared clutch size and 
weighted daily survival rates between experimental 
nests and nests of non-radioed females found by sys- 
tematic nest searches (Klett et al. 1986). We set deci- 
sion levels at & = 0.05. Our research was reviewed and 
approved (protocol 19920007) by the University of 
Saskatchewan Committee on Animal Care and Supply. 


Results 

We located 137 nests (64 control; 73 experimental). 
Experimental nests had a mean age when found of 
5.8 days (SD = 3.0). Fifty-nine of the 64 (92%) con- 


— eer 4 


2005 


trol nests were physically located after they were no 
longer active. Because their success or failure could not 
be determined and habitat class and vegetation type 
could not be verified, the five control nests not locat- 
ed were excluded from further analyses. Differences 
in hatching rates between nests where the female was 
flushed (19.2%; experimental) and those where she 
was not flushed (27.5%; control) approached signifi- 
cance (y* = 2.952, 1 df, P = 0.086). When 22 nests 
(30.1% of experimental nests) that were abandoned 
due to investigator activity were removed, hatching 
rates did not differ between the two groups (x7 = 0.294, 
1 df, P = 0.588). Mayfield estimates (Klett et al. 1986) 
of hatching rates for experimental and control groups 
were 19.9% (95% CI = 11.8 — 33.4%) and 20.8% 
(95% CI = 12.8 — 33.8%), respectively. Weighted 
daily survival rates of experimental (0.9537) and con- 
trol nests (0.9549) did not differ (F = 0.001, 1,5 df, 
P = 0.977). Nest loss due to predation occurred on 
average 10.1 days (SD = 7.8, n = 34) and 9.1 days 
(SD = 6.7, n = 35) following the date the nest was 
found or marked for experimental and control nests, 
respectively, and did not differ (t-test, t= 0.57, df = 67, 
P = 0.568). Weighted daily survival rates between 
experimental nests and nests of non-radioed females 
(0.9536, n = 44 nests) did not differ (F = 0.0002, 1,5 
df, P = 0.990). Mean clutch size between the two 
groups (8.8 eggs, SD = 1.5, n = 32 for experimental 
nests; 9.3 eggs, SD = 1.1, n = 34 for nests of non- 
radioed females) approached significance (f-test,’‘t = 
1.71, df = 64, p = 0.093). Using radio-telemetry alone, 
we correctly estimated the habitat class for 55 of 59 
(93%) control nests. Nest site vegetation was correctly 
estimated for 33 of 59 (56%) control nests. 


Discussion 

Low nesting success among prairie waterfowl (see 
Beauchamp et al. 1996), and particularly low nesting 
success among laying-stage nests (Miller and Johnson 
1978), often necessitate locating nests early in laying 
if certain reproductive data are to be collected. But 
protocols requiring female Mallards to be flushed from 
their nests early in laying increase the likelihood of 
nest abandonment. Our investigator-caused abandon- 
ment rate of 30.1% was high and similar to the 24.3% 
investigator-caused abandonment rate for radio-marked 
Mallards encountered during the first year (1993) of our 
study. Livezey (1980) reported an investigator-caused 
abandonment rate of 6% with most abandonment oc- 
curring among nests found prior to the fourth day of lay- 
ing. All investigator-caused nest abandonment (n = 22) 
in our study occurred the day the nest was first visit- 
ed. Nests had a mean age at abandonment of 4.2 days 
(SD =1.8). While younger-age nests are more prone to 
abandonment (Ducks Unlimited Canada, Institute for 
Wetland and Waterfowl Research, unpublished data), 
other, unknown factors also probably influence aban- 
donment, such as an individual female’s tolerance to 


THORN, EMERY, HOWERTER, DEVRIES, and JOYNT: INVESTIGATOR EFFECTS 


543 


disturbance. We suspect radio-transmitters had minimal 
impact on the likelihood that females would abandon 
their nests following disturbance, but cannot verify this. 
Weighted daily survival rates did not differ between 
experimental nests and nests of non-radioed females 
(P = 0.990). However, differences in mean clutch size 
approached significance (P = 0.093). We were unable 
to compare investigator-induced abandonment rates 
between experimental nests and nests of non-radioed 
females because in a traditional rotational nest-search- 
ing scheme (see Klett et al. 1986), where nests are mon- 
itored at intervals of 7-10 days, the number of nests that 
are abandoned (either due to investigator disturbance 
or other factors) may be under-estimated because pred- 
ators may destroy the nest before the next visit. 

In contrast to the high investigator-related abandon- 
ment rates that we observed, only 6 (3 experimental, 
3 control; 5.5%) nests were abandoned for reasons 
other than investigator disturbance. The mean age of 
abandonment for these nests was 12.5 days (SD = 6.2). 
Some of the highest reported incidences of natural 
abandonment were recorded by Duebbert et al. (1983) 
who found that Mallards nesting at high densities on 
an island abandoned 19%, 16% and 8% of their nests 
during the three years of their study. Because they did 
not visit the island during the nesting season, they 
concluded that inter- and intra-specific interactions, 
not investigator activity, were the major causes of aban- 
donment (Duebbert et al. 1983). 

When nests where investigator disturbance caused 
abandonment were removed from analysis, there was 
no difference in hatching rates or weighted daily sur- 
vival rates between experimental and control nests. The 
temporal pattern of nest loss also did not differ among 
groups. It would appear then, that flushing a female 
Mallard and visiting the nest once did not increase 
the likelihood of predation. Thirty-four of 51 (66.7%) 
experimental nests and 37 of 59 (62.7%) control nests 


_were destroyed by predators. Gotmark (1992) found 


a decrease in hatching rates attributable to investigator 
activity in 18% of published accounts where mammals 
were the dominant predators. In contrast, when avian 
predators dominated the egg-eating predator commu- 
nity, investigator disturbance led to increased preda- 
tion of eggs (Gotmark 1992). Both avian and mam- 
malian predators were common on our study sites 
(Ducks Unlimited Canada, Institute for Wetland and 
Waterfowl Research, unpublished data), yet we did not 
see increased predation resulting from investigators 
visiting nests once. 

Investigator activity during early egg-laying appears 
to impact Mallard hatching rates through a high rate 
of nest abandonment. Although most modern water- 
fowl studies account for nests for which the investi- 
gator caused abandonment when estimating hatching 
rates (Klett et al. 1986), investigator-caused abandon- 
ment may lead to increased renesting and, subsequent- 
ly, biased estimates of other reproductive parameters. 


544 


For example, re-nesting birds usually produce smaller 
clutches (Rohwer 1992). Also, late-hatched ducklings 
may have lower survival and recruitment rates (Dzus 
and Clark 1998; Dawson and Clark 2000; Anderson 
et al. 2001; Blums et al. 2002; and see Rohwer 1992 
for review of reproductive patterns). Causing birds to 
abandon their nests and, when re-nesting, select a new 
nest site also may bias estimates of habitat use and, if 
the new nest site has a different vulnerability to pre- 
dation than the original nest, hatching rates. Although 
investigator-induced abandonment rates in a tradition- 
al rotational nest-searching scheme (Klett et al. 1986) 
typically would not approach the rates found in this 
study because most of our nests were found early in 
laying, the extent to which traditional nest search stud- 
ies under-estimate abandonment rates should be inves- 
tigated further. Radio-telemetry should provide a use- 
ful tool for that investigation. 

Using radio-telemetry, we were able to collect accu- 
rate nesting information without disturbing the birds 
or visiting the nests until after the nests were no longer 
- active. We were able to locate over 90% of the nests 
after they had hatched, were destroyed, or were aban- 
doned by the female. Even if it had not been possible 
to locate such a high percentage of nests, we have 
demonstrated that it is possible to classify habitats with 
a high degree of accuracy using just telemetry. Not 
surprisingly, however, as the resolution of the habitat 
information became finer (i.e., nest site vegetation) 
our success rate declined. Nevertheless, radio-telemetry 
appears to provide a way to collect nesting data for 
waterfowl during the early egg-laying stage while avoid- 
ing the possibility of biasing data through increased 
abandonment rates. 

Study results led to protocol change for the remain- 
ing years (1995 — 2000) of the PHJV assessment pro- 
gram. We estimated nest locations via triangulation 
and physically located nests later when females were 
absent from the nest areas. Investigator-induced aban- 
donment due to accidental flushes during nest-marking 
averaged only 3.9% for the 22 study areas investigated 
during this period. 


Acknowledgments 

We gratefully acknowledge the cooperation of the 
Belmont, Davis and Erskine landowners for granting 
us access to their land. C. Archambault, B. Boldaun, 
D. Brooker, S. Cronin, F DeZeeuw, V. Fairman, M. 
Francis, K. Hento, N. Herbert, P. Johnston, T. Kro- 
likowski, D. Kuhn, G. Lacroix, C. Lamey, S. Leach, 
D. Marsh, K. Mott, A. Mullie, B. Parkin, K. Pilgrim, 
M. Post, K. Rattray, F. Rice, T. Sallows, D. Shaw, R. 
Terwilliger, P. Thorpe, J. Trevor, P. Varney, A. Viveiros, 
J. Walker and C. Wilke helped greatly with data col- 
lection. M. Barr, B. Calverley, K. Eskowich, A. Hak, 
M. Kornder, I. McFarlane, K. Remple, A. Richard, K. 
Schmitt and B. Thompson provided additional help 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


during decoy trapping and we express our thanks. 
Ducks Unlimited staff from Manitoba, Saskatchewan, 
and Alberta provided invaluable support during site 
selection and landowner contacts. We thank R. R. Cox, 
A. J. Erskine, and three anonymous reviewers for pro- 
viding helpful comments on earlier drafts of this man- 
uscript. Financial support for this study was provided 
by Ducks Unlimited Canada, Ducks Unlimited Inc., 
the Canadian Wildlife Service, the National Fish and 
Wildlife Foundation, and the North American Wetlands 
Conservation Council through the Institute for Wet- 
land and Waterfowl Research. 


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Received 3 August 2004 
Accepted 4 November 2005 


Food Habits of Dabbling Ducks During Fall Migration in a Prairie 
Pothole System, Heron Lake, Minnesota 


RYAN M. WERSAL!, Brock R. MCMILLAN?, and JOHN D. MADSEN! 


'GeoResources Institute, Mississippi State University, Box 9652, Mississippi State, Mississippi 39762, USA [corresponding 
authors | 

"Department of Biological Sciences, Minnesota State University, Mankato, 242 Trafton Science Center S, Mankato, Min- 
nesota 56001 USA 


Wersal, Ryan M., Brock R. McMillan, and John D. Madsen. 2005. Food habits of dabbling ducks during fall migration in a 
prairie pothole system, Heron Lake, Minnesota. Canadian Field-Naturalist 119(4): 546-550. 


We conducted an analysis of dabbling duck food habits in the fall of 2002 and 2003 in the Heron Lake system. Gizzard con- 
tents of hunter-harvested birds were analyzed using the percent aggregate volume method to determine what food items were 
consumed and in what quantity. Curltop Ladysthumb (Polygonum lapathifolium) was the food item consumed most often 
(82.2%) and in the greatest volume (34.2 ml). Sago Pondweed (Stuckenia pectinata) was the only food item of which multi- 
ple plant parts were consumed. However, the seeds and tubers only comprised 1.27 and 0.07 of the total aggregate percent. 


Key Words: Food habits, waterfowl, gizzard contents, aggregate volume, submersed macrophytes, Curltop Ladysthumb, 


Polygonum lapathifolium, Sago Pondweed, Stuckenia pectinata. 


The food habits of waterfowl are often diverse and 
vary throughout a state and flyway (Havera 1999). It is 
necessary to have a firm understanding of food use of 
waterfowl for effective management and to provide a 
diversity of high quality wetland habitat (Havera 1999). 
Quality habitat (abundant and available) is the most 
important ecological component affecting waterfowl 
populations which includes the essential elements of 
food, cover, and water (Baldassarre and Bolen 1994). 
Wetland habitats are crucial in providing food that wat- 
erfowl prefer instead of areas where they have to feed 
on only what is available (Havera 1999). Waterfowl con- 
sume a wide variety of vegetation of which submersed 
macrophytes constitute a large fraction of the total food 
items consumed (Martin and Uhler 1939; Havera 1999). 

Submergent macrophyte communities are a direct 
source of waterfowl food and indirectly serve as an 
environment for aquatic macroinvertebrates (Baldas- 
sarre and Bolen 1994). Submersed macrophytes can 
comprise the bulk of food items consumed by migrat- 
ing waterfowl (Martin and Uhler 1939). For example, 
Curlyleaf Pondweed (Potamogeton crispus) on aver- 
age yields 140 kg/ha of seed per season, or enough to 
sustain 2470 Mallards (Anas platyrhynchos) per hectare 
per day (Hunt and Lutz 1959). The pondweeds (Pota- 
mogeton and Stuckenia spp.) ranked first, by volume, 
as food consumed by 18 species of waterfowl (Martin 
and Uhler 1939). 

Of the pondweeds, S. pectinata is said to be one of 
the most sought after food plants by waterfowl (Kantrud 
1990). Stuckenia pectinata is probably the most impor- 
tant single waterfowl food plant on the continent and 
is responsible for about half, or more, of the total food 
percentage credited to the genus Potamogeton [Stuck- 


enia] (Martin and Uhler 1939). As a food item, S. pecti- 
nata can form a significant portion of foods found in 
gizzards of fall staging populations, pre-molting birds, 
flightless molting ducks, and ducklings (Chura 1961; 
Hay 1974; Keith and Stanislawski 1960). 

Historically, the Heron Lake system has been an im- 
portant staging area for fall migrating waterfowl due to 
its extensive beds of aquatic macrophytes. An account 
from 1906 reported over 700 000 Canvasbacks (Aythya 
valisineria) on North Heron Lake feeding on mats of 
Wild Celery (Vallisneria americana) (Berry and Ger- 
man 1999). The Wild Celery disappeared in 1922 as a 
result of the Common Carp (Cyprinus carpio) (Berry 
and German 1999) and today sago pondweed is the only 
submersed macrophyte found in the Heron Lake sys- 
tem (Case 2003; Case and Madsen 2004). Poor water 
quality is attributed to the lack of macrophyte growth, a 
likely result of runoff from the high percentage of agri- 
cultural land surrounding the lakes and point source pol- 
lution including sewage and industrial effluent from 
nearby cities (Case 2003; Case and Madsen 2004). The 
Heron Lake system also has high sediment resuspen- 
sion and increased turbidity due to shallow water and 
a large fetch (Case 2003; Case and Madsen 2004). 

Although the Heron Lake system is in a degraded 
state, it is still used by migrating waterfowl as it is one 
of the largest shallow lake systems in the region. Due to 
its continued importance to waterfowl we conducted a 
preliminary investigation of the food habits of fall 
migrating dabbling ducks staging in the Heron Lake 
system in October of 2002 and 2003. The primary ob- 
jective was to determine the principal foods utilized by 
fall migrating dabbling ducks, with particular interest 
to aquatic macrophytes, especially sago pondweed. 


546 


2005 


Methods 
Site Description 

The Heron Lake system is located in Jackson Coun- 
ty, Minnesota (43.72333°N 95.2325°W) and compris- 
es four lakes; of which South Heron and North Heron 
Lakes were used in this study. The four lakes have a 
mean depth of less than 1.5 m and a combined surface 
area of approximately 3200 hectares. South Heron Lake 
(1220 ha) can be divided into a north bay and a south 
bay by differing sediment characteristics (Case and 
Madsen 2004). South Heron Lake is connected to North 
Heron Lake via Division Creek located in the northern 
most part of the lake. North Heron Lake (1350 ha) is 
a flat, shallow lake with little change in depth from 
one end to the other. North Marsh (430 ha) is a small 
shallow water body located at the northern edge of 
North Heron Lake. Birds harvested on North Marsh 
were included in North Heron Lake due the close prox- 
imity of the lakes. Duck Lake (190 ha) was not included 
in this study due to the lack of public access for hunters. 


Food Habit Collection 

Hunters from South Heron and North Heron Lakes 
(including North Marsh) were asked to collect the giz- 
zards from their harvested birds during each hunting 
day in 2002 and 2003. The harvested samples were 
placed in 1.06-1 Whirl-pak bags filled with a 75% ethyl 
alcohol solution. A waterproof label was affixed to each 
bag to record the lake, the species and sex of the bird, 
and the date of harvest. The collected samples were 
stored in a refrigerator at approximately 4°C until pro- 
cessing. 

In the lab, each gizzard was opened and its contents 
washed through a series of standard testing sieves. 
Number 20 (850 micrometer), number 35 (500 micro- 
meter), and number 140 (106 micrometer) sieves were 
used to separate food items by size. Samples were pro- 
cessed using the dry volumetric method (Rogers and 
Korschgen 1966). Each food item was placed in plastic 
measuring dishes and dried at 55°C for 48 hours in a 
constant temperature oven. We summarized food items 
by species of waterfowl. Organic and inorganic materi- 
als were separated using a dissecting microscope. Items 
within a sample were separated according to species 
of plant or animal. 

Volumetric measurements were made using a micro- 
syringe (used for small food items and items found in 
small quantities), 10 ml, or 25 ml graduated cylinders 
(Rogers and Korschgen 1966). We placed food items 
into the micro-syringe or graduated cylinder and com- 
pressed slightly with a small dowel to remove any air 
pockets. Once the air had been removed we recorded 
the volume of that food. We identified plant items to 
genus level (if postmortem digestion had not ren- 
dered the item unidentifiable) and macroinvertebrates 
were identified to family. The identification of seeds 
was made using descriptions and diagrams by Martin 
(1951, 1954) (for Polygonum and Potamogeton spp.), 
the text by Martin and Barkley (1961), and the taxo- 


WERSAL, MCMILLAR, and MADSEN: FOOD OF DABBLING DUCKS 


547 


nomic key by Crow and Hellquist (2000). Seed iden- 
tifications were verified by a taxonomic expert at Mis- 
sissippi State University. Scientific and common names 
for plant species followed the Checklist of North 
American Plants for Wildlife Biologists by Scott and 
Wasser (1980). We identified invertebrates using the 
taxonomic key by Pennak (1978). 


Data Analysis 

We used percent occurrence to describe the frequen- 
cy of a food item in the samples and to assess the rel- 
ative use of a food item by waterfowl. We expressed 
volumes of food items as the percent of aggregate vol- 
ume (Martin et al. 1946). Aggregate volume quantifies 
the relative importance of food types as it addresses the 
amount of a food item consumed by waterfowl. Indi- 
viduals from the same species were aggregated per food 
item and the sum of each food item was then divided by 
the total volume of food consumed by all individuals. 

Percent Aggregate Volume was calculated as (1): 


> (volume of food item, across all animals in the sample) x 100 
Total volume of all foods consumed in the sample 


Results 
Harvested Waterfowl 

We collected 63 gizzards from fall migrating water- 
fowl, 46 gizzards in 2002 and 17 gizzards in 2003 
(Table 1). Blue-winged Teal (See Table 1 for scientific 
names) comprised the majority of the samples taken, 
followed by Green-winged Teal, Mallards, and Wood 
Ducks. Other species represented included the North- 
ern Shoveler and Northern Pintail. 

Plant material comprised 99.4% of the total aggre- 
gate percent of food items collected (Table 2). Polygon- 
um lapathifolium seeds made up the largest aggregate 
percent of food items collected from waterfowl in this 
wetland system. Potamogeton pusillus seeds ranked 
second in total aggregate percent followed by the seeds 
of Rice Cutgrass (Leersia oryzoides) and P. dichotomi- 
florum. Soybeans (Glycine max) appeared to rank high- 
er in aggregate percent than P. dichotomiflorum; how- 
ever, they were found in large amounts only in the few 
Wood Ducks harvested in this study. The fact that it 
was only found in a few individuals overestimated its 
volumetric importance. 

Animal matter constituted only (0.5%) of the total 
aggregate percent (Table 2). Water boatmen (Corixi- 
dae) were consumed more frequently and in the great- 
est amount. Blue-winged Teal accounted for the major- 
ity of the midges (Chironomidae) consumed. Other 
dabbling ducks that consumed invertebrates include 
Green-winged Teal, Mallards, and Gadwall which con- 
sumed Planorbidae, Physidae, and Plecoptera inverte- 
brates, respectively. 


Discussion 
Polygonum lapathifolium was the most important 
food consumed by dabbling ducks; it occurred in the 


548 THE CANADIAN FIELD-NATURALIST Vol. 119 


TABLE |. Species list of dabbling ducks harvested in the Heron Lake system during the fall hunting seasons of 2002 and 
2003. 


Species n % of Sample Female Male Unknown 
Blue-winged Teal (Anas discors) of 42.8 9 1 il 
Green-winged Teal (Anas crecca) 14 DR) Zz 4 3 
Mallard (Anas platyrhynchos) 13 20.6 8 5 0 
Wood Duck (Aix sponsa) 4 6.3 ! pu 1 
Northern Shoveler (Anas clypeata) 2 Bi l 1 0 
Gadwall (Anas strepera) 2 301 l 1 0 
Northern Pintail (Anas acuta) l 15 0 0) l 
Total 63 100.0 27 14 p) 


TABLE 2. Gizzard contents (+ | SE) of harvested dabbling ducks in the Heron Lake system during October of the 2002-2003 
hunting seasons. Food items without a standard error had less than two birds containing that food item. All food items are 
seeds unless otherwise noted. 


Food Item Frequency (%) Volume (ml) Aggregate (%) 
Amaranthus spp. 46.77 0.45 + 0.15 0.73 BO15 
Bidens spp. 9.68 0.05 + 0:02 0.08 + 0.01 
Carex spp. 40.32 0.24 + 0.06 0.39 + 0.06 
Ceratophyllum demersum 37.10 067-2 O18 1.09 + 0.18 
Chenopodium spp. 3.23 0.05 + 0.02 0.08 + 0.01 
Cyperus erythrorhizos 6.45 0.08 + 0.02 O13 O02 
Cyperus odoratus 46.77 0.34 + 0.03 0!55 40:08 
Echinochloa crusgalli 22.58 1.29 + 0.38 2.10) 0:38 
Eleocharis spp. 9.68 0.01 + 0.00 0.02 + 0.01 
Glycine max 1.61 2.80 4.57 
Leersia oryzoides 19:35 ZITO): 4.49 + 0.53 
Lolium spp. 1.61 0.03 0.05 
Panicum dichotomiflorum 41.94 2.24 + 0.88 3.65 + 0.88 
Pinus spp. 1.61 0.06 0.10 
Polygonum lapathifolium 82.26 2tOPE 173 34.27 + 1.73 
Polygonum pennsylvanicum 14.52 O15 :0,01 0.24 + 0.01 
Polygonum persicaria 8.06 0.02 + 0.01 0.03 + 0.01 
Potamogeton natans 8.06 0.37.+ (0:09 0.60 + 0.09 
Potamogeton pusillus 58.06 5.37 + 0.64 8.76 + 0.64 
Potamogeton (unidentified) 11.29 0.08 + 0.01 0.13 + 0.01 
Sagittaria latifolia 6.45 0.80 + 0.39 1.31 +0.39 
Scirpus acutus 41.94 Diy 0108 3:51 £0.08 
Scirpus fluviatilis 25.81 1.96 + 0.30 3.202040 
Scirpus validus 17.74 O17 = O:01 0.28 + 0.01 
Stuckenia filiformis 4.84 0.42 + 0.18 0.69 + 0.18 
Stuckenia pectinata 22-58 0.78 + 0.08 1.27 + 0.08 
Stuckenia pectinata (tuber) 1.61 0.04 0.07 
Zanichellia spp. 1.61 0.02 0.03 

Zea mays 1.61 6.80 11.09 
Setaria spp. 1.61 0.03 0.05 
Macerated Items 52.26 97S +035 15-91 O35 
Chironomidae 5.23 0.07 + 0.01 OPT O01 
Corixidae 16.16 OAD + O.02 0.20 + 0.02 
Physidae 1.61 0.07 0.11 
Planorbidae Bde) 0.01 + 0.00 0.02 + 0.01 
Plecoptera 1.61 0.02 0.03 
Unidentified Invertebrate 3323 0:03. 0:01 0.05 + 0.01 
Total Food 61.30 + 0.63 100.00 + 1.04 
Total Plant 60.98 + 0.75 99.48 + 1.22 


Total Animal 


0.32 + 0.01 


0.52 + 0.02 


2005 


majority of species, and ranked highest in percent ag- 
gregate volume. These results concur with a similar 
study conducted in Illinois where P. lapathifolium oc- 
curred in 95% of all Blue-winged Teal gizzards sam- 
pled (Havera 1999). Polygonum lapathifolium is wide- 
ly distributed in the Heron Lake system as noted by 
vegetation transects (Heron Lake Watershed District, 
unpublished data) making it readily available as a food 
source. Stuckenia pectinata was also readily available 
as a food source as it is the only submersed macrophyte 
found in the Heron Lake system (Case and Madsen 
2004). It had a frequency of occurrence of 74.5% and 
46.8% for North Heron and South Heron Lakes in 
2001 (Case and Madsen 2004). Likewise, in a similar 
study S. pectinata had a frequency of occurrence of 
77.6% and 52.8% for South Heron Lake in 2002 and 
2003, respectively; as well as 26.5% in North Heron 
Lake during 2003 (Wersal 2004). Stuckenia pectinata 
was the only item that dabbling ducks consumed mul- 
tiple parts of the plant. Seeds and tubers were found in 
gizzards analyzed in this study indicating that S. pecti- 
nata was utilized as a source of food for dabbling ducks. 
Kantrud (1986) estimated that a single S. pectinata 
dominated lake could support a large percentage of the 
continental migrating waterfowl population for a month 
during fall staging. 

The reduced abundance of §. pectinata observed in 
the Heron Lake system in 2003 may have shifted feed- 
ing to other areas of the system where other food items 
were more abundant. “Superior availability after all is 
the guiding principle in the choice of foods by birds” 
(McAtee 1918). Most dabbling species are non-selec- 
tive in their feeding habits and feed primarily on aquat- 
ic Or moist soil vegetation that is abundant in a given 
location (Havera 1999). Mallards have not been found 
to be selective foragers, but consume what is abundant 
in the area (Nummi 1993). Blue-winged Teal change 
their diets in relation to varying wetland conditions to 
take advantage of abundant food resources (Thomp- 
son et al. 1992). Northern Pintails have been found to 
shift their food selection based on availability of food 
items in a given area (Euliss and Harris 1987). 

Analyzing food habits in a given area using giz- 
zards as opposed to esophagus samples may introduce 
a bias toward hard-bodied food items as a result of the 
differences in breakdown rates of hard versus soft- 
bodied items (Swanson and Bartonek 1970). However, 
Wright (1959) found that Mallards contained approx- 
imately the same food items in their gizzards as they 
did in their esophagus. Furthermore, the esophagus of 
hunter-killed waterfowl are most often empty or con- 
tain small amounts of food (Drake 1970; Perry and 
Uhler 1982) resulting in the reliance on gizzards to 
obtain sufficient sample numbers (Havera 1999). Also, 
the bias of hard-bodied items may not be as critical in 
food habit analyses conducted in the fall because ani- 
mal matter is not as prevalent in the diets of most spe- 
cies of waterfowl, most notably dabbling ducks (Hav- 
era 1999). 


WERSAL, MCMILLAR, and MADSEN: FOOD OF DABBLING DUCKS 


549 


The dabbling ducks harvested in the Heron Lake sys- 
tem consumed a variety of food items during fall migra- 
tion. The most observed waterfowl species in the Heron 
Lake system were Mallards, Blue-winged Teal, Green- 
winged Teal, and Canada Geese, an observation support- 
ed by the gizzards collected in this study. Diving species 
still utilize the Heron Lake system; however, not to the 
extent of dabbling ducks. The decrease in use of the sys- 
tem by diving ducks can be attributed to the reductions 
in submersed macrophytes within the system. 

The results of this study suggest that submersed 
macrophytes were not as important to dabbling ducks 
as moist soil vegetation. Dabbling ducks are finding 
food; however, management should focus on providing 
more food that ducks prefer such as moist soil vegeta- 
tion instead of having ducks feed only on what is avail- 
able. Likewise, improvements in water quality should 
facilitate increases in S. pectinata populations and the 
re-establishment of other aquatic macrophytes to pro- 
vide a diversity of food items for diving species. Future 
studies in the system should focus attention towards 
collecting larger sample sizes; analyze changes in food 
availability based on management within the system; 
and analyzing esophagus samples to see if differences 
in food items exist from that of gizzards. 


Acknowledgments 

This project was funded by the U.S. Fish and Wild- 
life Service in cooperation with the Heron Lake Water- 
shed District and the North Heron Lake Game Produc- 
ers Association. A special thanks to Jason Neyens and 
Todd Boonstra for their diligent work in sorting through 
the gizzard samples. Victor Maddox provided verifica- 
tion of seed identifications. Earlier reviews of this man- 
uscript were provided by Eric Dibble, Victor Maddox, 
and Randy Markl. We would also like to thank those 
hunters who took the time to collect samples used in 
this study. Publication number J-10652 of the Missis- 
sippi Agriculture and Forestry Experiment Station. 


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Received 29 January 2005 
Accepted | November 2005 


Added in proof 

Wersal, R. M., J. D. Madsen, B. R. McMillan, P. D. Ger- 
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Body Weights of Adult and Juvenile Northern Pocket Gophers, 
Thomomys talpoides, in Central Alberta Alfalfa Fields 


GILBERT PROULX 


Alpha Wildlife Research & Management Ltd., 229 Lilac Terrace, Sherwood Park, Alberta T8H 1W3 Canada 


Proulx, Gilbert. 2005. Body weights of adult and juvenile Northern Pocket Gophers, Thomomys talpoides, in central Alberta 
alfalfa fields. Canadian Field-Naturalist 119(4): 551-559. 


In an effort to recognize adults from juveniles in late summer and early fall, carcasses of Northern Pocket Gophers (Thomomys 
talpoides) captured from April to October 1994 in a pure alfalfa (Medicago spp.) field in Camrose and a mixed alfalfa-orchard 
grass (Dactylis glomerata)-dandelion (Taraxacum spp.) field in Lacombe, in central Alberta were analyzed. Mean weights of 
adult males and females were significantly (P < 0.05) larger than those of juveniles with near-adult body sizes in late summer 
and early fall. On the basis of the limits of 95% confidence intervals for individual values in Camrose and Lacombe, live-captured 
males weighing < 130 g could be classified as juveniles; those that are > 180 g would be adults. Males weighing between 130 and 
180 g could be either juveniles or adults. Live-captured females weighing < 110 g would likely be juveniles. With larger animals, 
the pubic symphysis should be used to distinguish young females from adults. Given the variability of body weight in North- 
ern Pocket Gophers, it is suggested that criteria for aging based on weight be derived for local populations in specific habitat 
types. On average, Northern Pocket Gopher males and females from Camrose were significantly (P < 0.05) heavier than those 
of Lacombe. Animals from both fields were, on average, heavier than those previously studied in natural-vegetation commu- 


nities. Differences in the food nutritional quality may explain the observed variation in body weights among populations. 


Key Words: Northern Pocket Gopher, Thomomys talpoides, Alfalfa, Medicago sp., body weight, Alberta. 


In ecological studies, being able to recognize adults 
from juveniles in late summer and early fall without 
killing animals is useful to estimate summer mortality 
rates and population changes in response to environ- 
mental fluctuations (Caughley 1977), and conduct be- 
havioral and physiological investigations (Miller 1952). 
Northern Pocket Gopher (Thomomys talpoides) adults 
and juveniles are difficult to tell apart in late summer 
and early fall on the basis of weight. In British Colum- 
bia orchard-old field habitats, from July to September, 
Sullivan et al. (2001) used three age classes of North- 
ern Pocket Gophers based on body weight: juveniles 
(young of the year, which do not breed until the next 
year) < 49 g; subadults (individuals of which 50% are 
mature in the upper weight class) 49-60 g; and adults 
(2 50% of individuals sexually mature in the lowest 
weight class) = 61 g. However, previous studies on the 
body size of Thomomys talpoides (Smith 1940; Hansen 
1960; Hansen and Bear 1964; Tryon and Cunningham 
1968; Hansen and Reid 1973) and Thomomys bottae, 
Valley Pocket Gopher, (Howard and Childs 1959; Daly 
and Patton 1986; Patton and Brylski 1987) showed that 
animal weights varied among pocket gopher popula- 
tions due to habitat types, altitude, latitude, and seasons 
(Hansen and Reid 1973). Patton and Brylski (1987) 
found that growth rates and body sizes of pocket gophers 
inhabiting alfalfa monocultures were greater than those 
of pocket gophers inhabiting natural-vegetation com- 
munities. Then, one may wonder about the usefulness 
of aging criteria based on weight developed in different 
regions and habitats to study Northern Pocket Gopher 
populations in alfalfa (Medicago spp.) fields of central 
Alberta. 


According to Reid (1973), young-of-the-year North- 
ern Pocket Gophers would attain near-adult body 
weight approximately 180 days after birth. That is to 
say that, in Alberta alfalfa fields where young are born 
in April-May (Proulx 2002), they would be similar in 
body size to adults during August-October. However, 
the growth rate of juvenile pocket gophers may be high- 
ly variable from one population to another (Daly and 
Patton 1986), and Reid’s (1973) estimate may not be 
applicable to central Alberta populations. 

Few complete data have been published on body 
weights of Northern Pocket Gopher populations. Al- 
though the Northern Pocket Gopher is common in 
western Canada (Proulx 2004), little is known about 
body weights of age and sex classes from different 
habitats. Within the context of a multi-year ecological 
study of Northern Pocket Gophers in Alberta alfalfa 
fields, I collected carcasses to compare body weights 
of adults and juveniles during summer and early fall in 
order to determine weight classes that could be used to 
age animals in live-capture programs. I also wanted to 
compare the mean weights of adults from central Al- 
berta alfalfa fields to those reported in the scientific 
literature for other regions and habitat types. I hypoth- 
esized that in late summer-early fall: (1) on average, 
adults would be heavier than juveniles; (2) individual 
body weights of adults and juveniles would overlap; 
(3) body weight categories may be established to dif- 
ferentiate some juveniles from adults; and (4) mean 
body weights of adults inhabiting alfalfa fields would 
be larger than those of adults from natural-vegetation 
communities. 


Si 


552 


Methods 

The study was carried out in two alfalfa fields of 
central Alberta, in the communities of Camrose and 
Lacombe. The Camrose field was a 3-year-old pure 
alfalfa stand with a high forage yield and excellent 
second cutting (Alberta Agriculture 1992*). The La- 
combe field was approximately 5 years old, and cor- 
responded to a mixture of alfalfa with orchard grass 
(Dactylis glomerata) and abundant dandelion (Tarax- 
acum spp.). Alfalfa was cut once and thereafter the 
field was used as pasture. Spring adult populations 
were estimated at 22.4 animals/ha in Camrose (Proulx 
1997), and 23.5/ha in Lacombe (Proulx 1995*). The 
ratio male:female was 0.7:1 in Camrose, and 0.75:1 
in Lacombe. Mean litter size was significantly larger 
in Camrose (x = 6.4; standard deviation = 1.4) than 
in Lacombe (5.1 + 1.1) (Proulx 2002) 

In 1994, kill-trapping was conducted from 18 April 
to 27 October. Trapping methodology was described 
by Proulx (1997). Animals were individually marked 
in the field, and frozen. The weight, sex and age were 
determined in the laboratory. In central Alberta, fe- 
males produce only one litter (Proulx 2002). The size 
of the reproductive organs was used to distinguish 
young-of-the-year from adults. Young males have 
smaller testes, and young females have smaller ovaries 
and narrower uterine horns than adults (Hansen 1960; 
Hansen and Reid 1973). As precocial breeding is not 
known to occur in Alberta populations (Proulx 2002; 
Proulx and Cole 2002), the pubic symphysis was also 
used to distinguish young females from adults; in 
adult females the bones are resorbed, leaving a pubic 
gap (Hisaw 1924; Hansen 1960). 

An analysis of variance followed by the Tukey test 
was used to compare mean weights between Camrose 
and Lacombe population samples (Zar 1999). Student 
t-tests were used to compare mean body weights of 
Northern Pocket Gophers of Camrose and Lacombe 
populations to those of populations reported in the sci- 
entific literature. Probability values P < 0.05 were con- 
sidered statistically significant. 


Results 
Males 

Mean weights of Camrose adult males differed sig- 
nificantly (F, ,, = 5.9, P < 0.005) among samples col- 
lected from 18 April to 31 May. Animals captured in 
late May were significantly (P < 0.005) lighter than those 
collected earlier in the year (Table 1). In Lacombe, 
mean weights of adult males were similar (F, g, = 1.7, 
P > 0.05) among samples collected from 18 April to 
31 May (Table 1). 

Mean weights of juvenile males differed Eines 
ly among samples in Camrose (F, 7, = 21.7, P < 0.005) 
and Lacombe (F,,¢, = 11.3, P < 0.005) (Table 1). 
Juveniles captured in September-October were the 
heaviest; they were similar in size (P > 0.05) to those 
captured in August-September (Table 1). 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Mean weights of adults and juveniles differed sig- 
nificantly in Camrose (F,7, = 21.7, P < 0.005) and 
Lacombe (F; ,;, = 80.2, P < 0.005). In both study 
areas, the mean adult weight was larger (P < 0.001) 
than that of each juvenile sample collected from June 
to October (Table 1). In Camrose, the limits of 95% 
confidence intervals for individual values were 142- 
230 g for adults, and 108-172 g for September-October 
juveniles. In late summer-early fall, although males 
were, on average, heavier than juveniles, there was a 
marked overlap between age class weights. Then, on 
the basis of weight alone, live-captured animals weigh- 
ing between 142 and 172 g could be classified as either 
adults or juveniles. Those weighing < 142 g could be 
juveniles while those > 172 g could be adults. In La- 
combe, the limits of 95% confidence intervals for indi- 
vidual values were 136-213 g for adults, and 106-174 g 
for September-October juveniles. Live-captured ani- 
mals weighing between 136 and 174 g could then be 
classified as either adults or juveniles. Those weighing 
< 136 g could be juveniles while those > 174 g could 
be adults. 


Females 

In Camrose, mean weights of adult females were 
similar (F; ,,3 = 1.0, P > 0.05) among samples collect- 
ed from 18 April to 27 June (Table 1). In Lacombe, 
mean weights of adult females differed significantly 
(Fy 197 = 3.0, P < 0.05) among samples collected from 
18 April to 27 October. Animals captured in late May 
and in August-October were the only samples to differ 
significantly (P < 0.005) from each other (Table 1). 

Mean weights of juvenile females differed signifi- 
cantly among samples in Camrose (F,3, = 20.3, 
P < 0.005) and Lacombe (F; 3, = 11.8, P < 0.005) 
(Table 1). In Camrose, juveniles captured in August- 
September and in July were of similar (P > 0.05) size, 
but were larger (P < 0.05) than those of June (Table 
1). In Lacombe, juveniles captured from August to 
October were of similar (P > 0.05) size, but were 
larger (P < 0.05) than those of July (Table 1). 

Mean weights of adults and juveniles differed signif- 
icantly in Camrose (F; 154 = 120.5, P < 0.005) and in 
Lacombe (Fy ,45 = 27.5, P < 0.005). In both study areas, 
the mean weight of adults was larger (P < 0.001) 
than that of each juvenile sample collected from June 
to September (Table 1). In Camrose, the limits of 95% 
confidence intervals for individual values were 126- 
195 g for adults, and 89-163 g for August-September 
juveniles. Although female adults were, on average, 
heavier than juveniles, there was a marked overlap 
between age class weights. Therefore, on the basis of 
weight alone, live-captured animals weighing between 
126 and 163 g could be either adults or juveniles. 
Females weighing < 126 g could be juveniles while 
those > 163 g could be adults. In Lacombe, the limits 
of 95% confidence intervals for individual values were 
113-175 g for adults and 101-180 g for September- 


Bb, 


Bopby WEIGHTS OF NORTHERN POCKET GOPHERS 


PROULX 


2005 


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554 


October juveniles. Live-captured animals weighing 
< 113 g could be classified as juveniles; those weigh- 
ing 2 113 g could be either adults or juveniles. 


Comparison between adult populations 

Mean weights of Camrose and Lacombe adults 
differed significantly (F; 3,,; = 86.7, P < 0.005). On 
average, Camrose males and females were heavier 
(P < 0.005) than those of Lacombe. 

On average, adult males from Camrose and Lacombe 
were significantly (P < 0.05) heavier than adult males 
from other alfalfa fields or habitat types (Table 2). On 
average, adult females from Camrose were also sig- 
nificantly (P < 0.05) heavier than females from other 
habitat types and regions. The mean weight of Lacombe 
adult females was, however, similar (P > 0.05) to those 
of females inhabiting alfalfa fields in Saskatchewan 
and Colorado (May sample), and unspecified agricul- 
tural lands in central Alberta (Table 2). 

The Camrose and Lacombe adult males and females 
were at least 25% heavier than males and females found 
in habitats other than alfalfa fields. They were more 
than twice as heavy as Northern Pocket Gophers cap- 
tured in British Columbia (Table 2). 


Discussion 

In this study, the hypothesis that adults would be, on 
average, heavier than juveniles in late summer-early 
fall was validated. Adult females grow until 2 years old, 
and males continue to grow throughout their lifetime 
(Tryon 1947; Miller 1952; Howard and Childs 1959; 
Daly and Patton 1986). Even though young-of-the-year 
appeared as large as adults, they were approximately 
6-7 months old in the fall, and their growth was not 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


completed (Tryon 1947). On the other hand, body 
sizes of juveniles and adults vary considerably within 
the same habitat due to differential individual growth 
rates from a same litter or population (Tryon 1947; 
Daly and Patton 1986), age of the individuals (i.e., older 
juvenile vs. younger adult) (Hansen and Bear 1964), 
and time of year (Miller 1952; this study). All this may 
explain the marked overlap observed within juvenile 
populations from August to October, and between 
juveniles and adults, as reported in previous studies 
(Tryon 1947; Miller 1952; Hansen and Bear 1964). 
This study, and previous work (Tryon 1947; Hansen 
and Reid 1973; Hansen and Bear 1964), showed that 
mean body weights of age and sex classes varied con- 
siderably within and between study areas. Therefore, 
it is unlikely that reliable body weight categories may 
be established to differentiate juveniles from adults. At 
best, confidence intervals for individual values may be 
cautiously used to recognize lighter juveniles and heav- 
ier adults. Using confidence intervals of individual 
values for juvenile and adult males in Camrose and 
Lacombe, I propose that, in late summer-early fall, 
live-captured animals weighing < 130 g would likely 
be juveniles. Individuals weighing > 180 g may be 
adults. Animals weighing between 130 and 180 g might 
be either juveniles or adults. In the case of females, 
animals weighing <110 g would likely be juveniles. 
Larger animals should be classified on the basis of the 
presence of an open pelvic girdle, i.e., the pubic bones 
are widely separated due to the permanent resorption 
of the symphysis at first pregnancy (Miller 1952). 
Given the variability of body weight in pocket 
gophers, any criteria for aging based on weight will 


TABLE 2. Mean weights of adult Northern Pocket Gophers reported in the scientific literature. 


Adult weights (g) 


Males Females 
Standard Standard 
Location Habitat types Period n xX deviation mm *X deviation 
Alberta Unspecified agricultural land near 
Edmonton — MacDonald (1969) Summer 43; 133 14 37. 138 20 
British Forb-orchard — Sullivan et al. (2001) Summer 1982 23 69 2 23 68 Z 
Columbia Old field-orchard — Sullivan et al. (2001) Summer 1983 oo Ti 2 49 60 | 
Orchard — Sullivan et al. (2001) Summer 1983 22 61 4 Do” 103 | 
Summer 1985 22)° TA 4 59° ‘65 Z 
Colorado Alfalfa field — Hansen (1960) April 86 159 61 49 136 20 
May 58). 558 15 36: 135%, 20 
June 18 148 21 25; 136. 16 
August © 158 19 D2) NS55 sao 
September o 176 17 12 1358" 5 
Alpine tundra — Hansen and Bear (1964) August 14 114 8 84. 306) 2 
Meadow — Hansen and Bear (1964) August 8 134 18 298 TORS AZ 
Shrub-bunchgrass — Hansen and Bear (1964) August 8 116 12 30° 105 12 
Weedy rangeland — Hansen and Ward (1965) August ay -A2Y 13 46 114 14 
Saskatchewan Alfalfa field — Runnells (1988) Summers 1983 
and 1984 32. 138 41 Ti, 4d 6S 
Utah Coniferous forest — Andersen (1978) June-October 5 104 3 14 91 2 


2005 


have to be derived for local populations in specific 
habitat types. Body weight confidence intervals deter- 
mined in this study may be useful to researchers work- 
ing on Northern Pocket Gophers in central Alberta 
alfalfa fields. These confidence intervals would not be 
reliable to age live-captured animals from meadows, 
old field or orchard populations. Habitat differences 
in nutritional quality of available food can directly 
affect the observed variation in body weight among 
populations of pocket gophers (Smith and Patton 1980; 
Patton and Brylski 1987). Alfalfa is a superior forage 
producing more protein per ha than any other crop or 
grass (Heath et al. 1973). Northern Pocket Gophers 
inhabiting Camrose and Lacombe alfalfa fields were 
heavier than animals from other habitat types. A dif- 
ference in the quality of food available in pure and 
mixed alfalfa fields may also explain the difference in 
the mean weights of Northern Pocket Gophers inhab- 
iting Camrose and Lacombe. 


Acknowledgments 

I thank Pam Cole, Joel Nicholson, Nicole Proulx, 
Orest Litwin and Paul King for technical support, and 
Pauline Feldstein for reviewing an earlier draft. I am 
especially grateful to referee David Nagorsen for pro- 
viding an insightful review of an earlier draft of this 


paper. 


Documents Cited (marked * in text) 

Alberta Agriculture. 1992. Alberta forage manual. Agdex 
120/20-4, Edmonton, Alberta. 

Proulx, G. 1995. Development of a northern pocket gopher 
border control strategy. Alpha Wildlife Research and Man- 
agement Ltd. report prepared for the Counties’ Pocket 
Gopher Control Program, County of Red Deer, Alberta. 


Literature Cited 

Andersen, D. C. 1978. Observations on reproduction, growth, 
and behavior of the northern pocket gopher (Thomomys 
talpoides). Journal of Mammalogy 59: 418-422. 

Caughley, G. 1977. Analysis of vertebrate populations. John 
Wiley & Sons Ltd., New York New York. 234 pages. 

Daly, J. C., and J. L. Patton. 1986. Growth, reproduction, and 
sexual dimorphism in Thomomys bottae pocket gophers. 
Journal of Mammalogy 67: 256-265. 

Hansen, R. M. 1960. Age and reproductive characteristics of 
mountain pocket gophers in Colorado. Journal of Mam- 
malogy 41: 323-335. 

Hansen, R. M., and G. D. Bear. 1964. Comparison of pock- 
et gophers from alpine, sub-alpine, and shrub-grassland 
habitats. Journal of Mammalogy 45: 638-640. 

Hansen, R. M., and V. H. Reid. 1973. Distribution and adap- 
tations of pocket gophers. Pages 1-19 in Pocket gophers 
and Colorado mountain rangeland. Edited by G. T. Turner, 
R. M. Hansen, V. H. Reid, H. P. Tietjen, and A. L. Ward. 
Colorado State University, Fort Collins, Bulletin 5548S. 
90 pages. 

Hansen, R. M., and A. L. Ward. 1965. Some relations of 
pocket gophers to rangelands on Grand Mesa, Colorado. 


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255 


Colorado Agricultural Experiment Station, Technical Bul- 
letin 88. 

Heath, M., D. S. Metcalfe, and R. F. Barnes. 1973. Forages. 
The science of grassland agriculture. Third edition. The 
Iowa State University Press, Ames, Iowa. 755 pages. 

Hisaw, F. L. 1924. The absorption of the pubic symphysis of 
the pocket gopher, Geomys bursarius (Shaw). American 
Naturalist 58: 93-96. 

Howard, W. E., and H. E. Childs, Jr. 1959. Ecology of 
pocket gophers with emphasis on Thomomys bottae mewa. 
Hilgardia 29: 277-358. 

MacDonald, S. J. 1969. A taxonomic study of the pocket 
gophers of Alberta. Mammalia: Geomyidae: Thomomys 
talpoides (Richardson). M.Sc. thesis, University of Alberta, 
Edmonton. 70 pages. 

Miller, M. A. 1952. Size characteristics of the Sacramento 
Valley pocket gopher (Thomomys bottae navus Merriam). 
Journal of Mammalogy 33: 442-456. 

Patton, J. L., and P. V. Brylski. 1987. Pocket gophers in alfal- 
fa fields: causes and consequences of habitat-related body 
size variation. The American Naturalist 130: 493-506. 

Proulx, G. 1997. A northern pocket gopher (Thomomys tal- 
poides) border control strategy: promising approach. Crop 
Protection 16: 279-284. 

Proulx, G. 2002. Reproductive characteristics of northern 
pocket gophers, Thomomys talpoides, in Alberta alfalfa 
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Proulx, G. 2004. Effects of female scents on the trappability 
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Protection 23: 1055-1060. 

Proulx, G., and P. J. Cole. 2002. Evidence of a second litter 
in northern pocket gophers, Thomomys talpoides. Canadian 
Field-Naturalist 116: 322-323. 

Reid, V. H. 1973. Population biology of the northern pocket 
gopher. Pages 21-41 in Pocket gophers and Colorado moun- 
tain rangeland. Edited by G. T. Turner, R. M. Hansen, V. 
H. Reid, H. P. Tietjen, and A. L. Ward. Colorado State 
University, Fort Collins, Bulletin 5548S. 90 pages. 

Runnells, J. 1988. Some aspects of population dynamics with- 
in a southern Saskatchewan population of northern pocket 
gophers (Thomomys talpoides). M.Sc. thesis, University 
of Regina, Saskatchewan. 116 pages. 

Smith, C. F. 1940. Weights of pocket gophers. Journal of 
Mammalogy 21: 220. 

Smith, M. F, and J. L. Patton. 1980. Relationships of pocket 
gophers (Thomomys bottae) of the lower Colorado River. 
Journal of Mammalogy 61: 733-756. 

Sullivan, T. P., D. S. Sullivan, and E. J. Hogue. 2001. Re- 
invasion dynamics of northern pocket gopher (Thomomys 
talpoides) populations in removal areas. Crop Protection 
20: 189-198. 

Tryon, C. A., Jr. 1947. The biology of the pocket gopher 
(Thomomys talpoides) in Montana. Montana State Col- 
lege Agricultural Experimental Station, Technical Bulletin 
448. 30 pages. 

Tryon, C. A., and H. N. Cunningham. 1968. Characteristics 
of pocket gophers along an altitude transect. Journal of 
Mammalogy 49: 699-705. 

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tice-Hall, Inc. Upper Saddle River, New Jersey. 663 pages. 


Received 30 November 2004 
Accepted 6 October 2005 


Protocole de suivi des populations d’aster du Saint-Laurent, 
Symphyotrichum laurentianum, aux Iles-de-la-Madeleine 


GUILLAUME de LAFONTAINE 


Département de biologie, Centre d’études nordiques, Université Laval, Québec, Québec G1K 7P4 Canada; Courriel: guil- 
laume.de-lafontaine. 1 @ulaval.ca 


de Lafontaine, Guillaume. 2005. Protocole de suivi des populations d’aster du Saint-Laurent, Symphyotrichum laurentianum, 
aux Iles-de-la-Madeleine. Canadian Field-Naturalist 119(4): 556-568. 


Laster du Saint-Laurent (Symphyotrichum laurentianum (Fernald) Nesom) est une plante endemique 2 a la région du golfe du 
Saint-Laurent et on ne la retrouve qu’au Nouveau-Brunswick, a I’[le du Prince-Edouard ainsi qu’ aux Iles-de-la-Madeleine, au 
Québec. Les populations les plus nombreuses se retrouvent aux Iles-de-la-Madeleine. L’ espéce est considérée «préoccupante» 
par le COSEPAC depuis avril 1989 puis menacée depuis 2004. De nombreuses études réalisées par des équipes distinctes depuis 
1988 ont tenté de recenser |’ effectif et la superficie des populations réparties sur I’ aire de distribution. Ces études étant réalisées 
par différentes équipes utilisant des méthodes distinctes pour mener les évaluations ont conduit a des résultats contrastants. 
Une revisite des occurrences connues aux Iles-de-la-Madeleine en septembre 2004 visait 4 mettre a jour I’ état des connaissances 
sur la localisation exacte et l’effectif des populations d’aster sur l’archipel. Aussi, un protocole de suivi simple a été élaboré 
sur place, a partir duquel le personnel de Il’ organisme Attention Frag’ [les pourra récolter les données dans le futur. Ces don- 
nées pourront ensuite étre cumulées et interprétées facilement par ministére développement durable environnement et parcs 
du Québec pour effectuer le suivi a long terme. La méthodologie, ainsi uniformisée, permettra de décrire les variations tem- 
porelles de l’état des populations en s’assurant qu’elles ne sont pas obscurcies par différentes méthodes de prise de données. 


Mots clés : Aster du Saint-Laurent, Aster laurentianus, Symphyotrichum laurentianum, espece endémique, espéce menacée, 
espéce rare, espéce vulnérable, golfe du Saint-Laurent, Iles-de-la-Madeleine, suivi de populations. 


Laster du Saint-Laurent est une plante endémique ment stables et fluctuaient peu d’une année a |’autre 
de la région du golfe du Saint-Laurent. On la retrouve en absence de perturbations anthropiques (Houle et 
au Nouveau-Brunswick, a I’fle du Prince-Edouard et —_ al. 2002). Enfin, suite & un relevé effectué en 2003, 
au Québec autour des lagunes sur l’archipel des [les- | Duclos notait que l’aster était en déclin sur la totalité 
de-la-Madeleine. Puisque son aire de distribution est de son aire de répartition (Duclos 2004). 
trés limitée et qu’elle n’est répartie qu’en petites popu- Ainsi, aux Iles-de-la-Madeleine, ot I’on retrouve les 
lations, ’'espece a d’abord été considérée comme _ plus grands effectifs, il y a eu quatre relevés depuis 
«préoccupante» par le COSEPAC en avril 1989, puis 1983. Ces quatre relevés menés par quatre équipes dis- 
son statut a été réexaminé en mai 2004 pour la désigner _tinctes présentent des résultats contrastant que les 
«menacée» (COSEPAC 2004). Ainsi, quoique décrite auteurs interprétent différemment. Certains concluent 
pour la premiére fois en 1914 par Fernald, les études que les populations fluctuent beaucoup (Gilbert et al. 
sur la localisation et l’effectif des différentes popula- 1999; Duclos 2004), alors que d’ autres, au contraire, 
tions n’ont véritablement débuté qu’en 1988. Depuis, concluent que les populations sont dans un état stable 
différentes équipes de recherche ont étudié la situation (Houle et al. 2002). On note cependant plusieurs lacunes 
démographique de |’ aster. concernant les méthodologies d’échantillonnage au 

Un premier rapport sur le statut de l’aster du Saint- sein de toutes ces études. Ainsi, ce qui constitue une 
Laurent au Canada a établi les populations et leurs population pour I’un est divisé en de nombreuses popu- 
effectifs estimés entre 1983 et 1986 (Houle and Haber __ lations pour l’autre et chacun utilise sa propre numéro- 
1990). Les informations ultérieures proviennent de _ tation pour désigner des populations définies de fagon 
relevés effectués 4 I’Ie du Prince-Edouard en 1992 et —_ arbitraire. Aussi, les visites sur le terrain ont été effec- 
1993 (Guignion et al. 1995) et aux Iles-de-la-Made- _ tuées a différents moments durant la saison de crois- 
leine en 1994 et 1995 (Gagnon et al. 1995a, 1995b). — sance. Sachant que I’aster est une espéce annuelle, la 
A partir de la comparaison des résultats des études _ phénologie peut donc s’avérer d’une importance capi- 
mentionnées ci-haut, Gilbert et al. (1999) soulignaient tale pour l’estimation de la taille des populations. Pour 
une fluctuation importante de la taille des populations _définir !’emplacement des populations, certains util- 
dans le temps et attribuaient cette variation a la dy- _isent des photos aériennes, une méthode rapide et pra- 
namique des perturbations naturelles ou anthropiques __ tique mais inexacte, alors que d’autres utilisent une 
de habitat. Or, un troisiéme relevé effectué aux Iles- seule coordonnée géoréférencée pour indiquer une pop- 
de-la-Madeleine entre 1999 et 2001 concluait que les ulation pouvant couvrir une superficie de plusieurs kilo- 
populations d’aster du Saint-Laurent étaient relative- | métres. Dans d’ autres cas, la seule information est un 


Nn 


56 


2005 


point sur une carte grossiere. Par conséquent, la locali- 

sation et l’effectif des populations demeurent équivo- 

ques. 

La présente étude avait pour but d’uniformiser la 
récolte de l’information sur |’aster du Saint-Laurent, 
afin de permettre au ministére du développement dura- 
ble, environment et parcs du Québec de suivre a long 
terme les variations temporelles de |’état des popula- 
tions en s’assurant qu’elles ne sont pas obscurcies par 
différentes méthodes de prise de données. L’ organisme 
local Attention Frag’Iles a été mandaté par le min- 
isteére du développement durable, environment et parcs 
afin d’assurer la collecte annuelle des données aux 
Iles-de-la-Madeleine. 

Les objectifs spécifiques de la présente étude sont : 
(1) Mettre a jour les données sur les effectifs, les 
localisations et les superficies des populations d’ aster 
du Saint-Laurent aux Iles-de-la-Madeleine en 2004. 
(2) Elaborer un protocole d’échantillonnage sim- 
ple afin de permettre un suivi a long terme. Ce pro- 
tocole a été remis au personnel d’ Attention Frag’ Iles 
basé aux Iles-de-la-Madeleine qui effectuera la ré- 
colte des données annuellement. Les données seront 
remises au ministere de du développement durable, 
environment et parcs du Québec qui assurera le 
suivi a long terme. 

(3) Décrire la situation actuelle des populations sé- 

lectionnées pour le suivi. 


Espece 

Symphyotrichum laurentianum (Fernald) Nesom 
est une plante halophyte annuelle, pratiquement glabre 
et dont la taille varie de 2 a 50 cm de hauteur. La tige 
est simple lorsque la densité est élevée ou ramifiée 
lorsque la densité diminue (observation personnelle). 
Les feuilles sont sessiles et entieres, parfois un peu 
charnues. Les fleurs sont regroupées en capitules en- 
tourés de bractées foliacées. Un capitule est présent a 
chaque ramification de la tige. Les premiéres plan- 
tules émergent a la fin de mai et les premieres inflo- 
rescences apparaissent au début du mois d’aoit alors 
que les fruits (akenes) matures sont disséminés par le 
vent vers la fin de septembre. 

Aux Iles-de-la-Madeleine, lespéce se retrouve typi- 
quement sur une bande de ca. 50 a 200 cm de largeur 
en périphérie des lagunes et des marais salés (Houle et 
al. 2001). Cette répartition limitée est due 4 une dyna- 
mique particuliére. Prés de la rive, l’émergence de 
Paster est limitée par la salinité (Houle et al. 2001, 
Reynolds et al. 2001; Reynolds et Houle 2002). La 
distribution prés du rivage est également affectée par 
exposition aux vents, aux vagues lors de tempétes, aux 
fortes marées d’équinoxe et aux dépéts de débris, sur- 
tout constitués de zostére (Zostera marina L.). Ce gra- 
dient de salinité et de perturbations physiques diminue 
en s’éloignant du rivage, favorisant ainsi la croissance 
de différentes especes. Il a de plus été démontré ex- 
périmentalement que le taux de croissance relatif de 


DE LAFONTAINE: L’ ASTER DU SAINT LAURENT AUX ILES-DE-LA-MADELEINE 


537 


Paster du Saint-Laurent diminue lorsque la plante est 
placée a l’ombre (Reynolds et al. 2001; Houle et 
Valéry 2003). Ainsi, l’aster est rapidement remplacé 
par d’autres especes plus compétitrices lorsque les 
conditions permettent la croissance de ces derniéres 
(Brumbt 2001). Il s’agit donc d’une espéce tolérante a 
un certain niveau de stress (salinité, inondations, dépdts 
de zostére, vents) qui restreint les autres especes plus 
compétitrices (Grime 1977, 1979). 


Méthodes 
Mise a jour des données sur les effectifs, les localisa- 
tions et les superficies des populations 

Les 25 stations de Houle et al. (2002) ou l’aster était 
présent (entre 1999 et 2001) et une station recensée 
par Gagnon et al. (1995b) mais non visitée par Houle 
et al. ont été revisitées du 8 au 16 septembre 2004, 
soit pendant la période de floraison / fructification. 
Ce relevé inclut donc toutes les dernieres populations 
recensées, mais exclut les occurrences historiques. 
Les populations ont été renommées selon des critéres 
géographiques plutot que numérotées arbitrairement 
comme dans plusieurs études précédentes. Ceci a pour 
but d’uniformiser la nomenclature et de classer les 
populations selon un critére fixe plut6t que sur une 
base potentiellement variable tel que la distance entre 
deux occurrences. L’effectif de chaque population a 
d’abord été estimé visuellement et une classe sur une 
échelle logarithmique de 0 a 6 a été attribuée a la popu- 
lation (classes : 0 aucun individu retrouvé; / — 1 a 10 
individus; 2 — 10 a 100 individus; 3 — 10? a 10° indi- 
vidus; 4 — 10° a 10* individus; 5 — 10* a 10° individus; 
6 — 10° a 10° individus). La localisation exacte de I’en- 
semble de la superficie couverte par chaque popula- 
tion a été définie par des coordonnées géoréférencées 
a l’aide d’un GPS. 


Elaboration d’un protocole de suivi simple 

Les trois populations ayant les effectifs les plus im- 
portants ont été sélectionnées aux fins du programme 
de suivi. Pour chacune des trois populations, cing tran- 
sects permanents ont été installés perpendiculaire- 
ment a la ligne du rivage. Les cing transects ont été 
répartis dans |l’ensemble de la population de fagon a 
couvrir l’ensemble des conditions environnementales 
permettant la croissance de |’aster. Tous les transects 
ont une dimension de 3 m de longueur par | m et sont 
divisés en six quadrats de 0.5 m par | m. Ainsi, chaque 
transect couvre la largeur de la bande d’ aster typique- 
ment retrouvée le long des lagunes. Le premier quadrat 
se trouve pres de la ligne d’eau dans le milieu plus 
perturbé et le dernier quadrat est situé a travers la végé- 
tation, dans le milieu plus compétitif. 

Les mesures devront étre prises annuellement, entre 
la deuxieme semaine de aoit et la troisieme semaine de 
septembre, soit pendant la période de floraison de I’as- 
ter. Dans chaque quadrat, huit variables importantes 
devront étre mesurées : (1) le nombre de tiges d’ aster; 
(2) le pourcentage de couverture par I’ aster; (3) le pour- 


558 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLEAU 1. Liste des occurrences d’aster du Saint-Laurent aux Iles-de-la-Madeleine en 2004 en comparaisons avec les 
effectifs observés dans les études précédentes. Les nombres entre parenthéses a droite du nom des populations référent aux 
stations de Houle et al. (2002). Les chiffres en caractéres gras indiquent la classe attribuée a la population dans la présente 


étude. 


Nom de la population 
Présente étude 


Cap de I’h6pital (11 et 12) 4- 1034 10* 
Barachois, Chemin Poirier (8 et 10) L= Lalo 
Barachois (9) 5- 10°74 106 
Etang du nord (13) 0 - aucun aster 
Havre aux Basques (14) 5- 1074 10° 
Pointe des véliplanchistes (15, 16 et 17) 3- 107410° 
Pointe aux canots (18) 3- 1074 10° 
Dune de |’ ouest (19) 3- 1074 103 
Etang des caps (20) 6- 10°a 10° 
Baie du portage 1- 1a 10 
Baie de Grosse Ile (43 et 44) 5- 10*a10° 
Baie Clarke (31) 4:2 10°'4 10* 
Pointe Old Harry (6) 0 - aucun aster 
Bassin aux Huitres, Bassin est (4 et 5) 3- 1074 10° 
Bassin aux Huitres, Club vacance (34) 2- 10a 100 
Bassin aux Huitres, centre (33) Non revisitée 
Bassin aux Huitres, Bassin ouest (2 et3) 3- 1074 10° 


Pointe de |’ est (23 et 24) Non revisitée 


centage de couverture par toute la végétation; (4) le 
pourcentage de couverture des dépéts de zostére; (5) 
le pourcentage de couverture de sol nu; (6) la hauteur 
de la plus haute tige d’ aster; (7) le nombre de capitules 
de la tige d’aster la plus ramifiée; (8) la richesse spé- 
cifique. Ces variables sont faciles a récolter, ne néces- 
sitent aucun instrument de mesure autre qu’un ruban 
a mesurer et sont rapidement accessibles. Cependant, 
les mesures devront idéalement étre effectuées par un 
observateur unique puisque les variables de recou- 
vrement sont évaluées par estimation visuelle et sont 
donc sujettes a changer selon |’ observateur. Les vari- 
ables sélectionnées donnent un indice de l’effectif, la 
densité, |’habitat et la morphologie de la population a 
étude. 


Description de la situation actuelle aux populations 
sélectionnées pour le suivi 

Les mesures aux trois populations ont été prises 
selon la méthode décrite a l’objectif précédent entre le 
16 et le 21 septembre 2004. Les analyses statistiques 
ont été effectuées a l’aide du logiciel Statistica release 
5 (StatSoft, Tulsa, Oklahoma, USA). II est 4 noter que 
les variables pourcentage de couverture du dépdt de 
zostere et pourcentage de couverture de sol nu n’ont 
pas été utilisées dans les analyses. Ces variables four- 
nissent un indice de l’habitat, mais ne seront pas dis- 
cutées en détail. 

Une analyse de variance nichée de modele I pur 
(pure Model II nested ANOVA) ot le facteur transect 
est niché dans le facteur population a été effectuée 
(Sokal et Rohlf 1969). Ceci permet de déceler 1’ exis- 
tence de véritables différences entre les populations 
d’une part et les cing transects de chaque population 


Nombre d’individus 


Duclos (2004) Houle et al. (2002) Gilbert et al. (1999) 
Ogee be 1074 10° 10*a 10° 
10°a 10° 1 a 1000 

10°4 10° 

Non revisitée 1 a 1000 10° 4 10° 
10*a 10° + de 10° 

Non revisitée 10* 4 10° 
107 a 10° 10°a 10° 

Non revisitée 10°4 10’ 

Non revisitée 10°a 10’ 

Non revisitée Non revisitée 

Non revisitée 107107 Non connue 

Non revisitée 10° a 10* 10.4 100 

Non revisitée 10a 100 0 a 1000 

0 10? 4 10° 10°a 10° 

0 10 a 100 

0 1a 100 

1034 104 10! 4 10° 
Non revisitée 10a 100 10 a 100 


d’autre part. Pour toutes les variables générant des ré- 
sultats significatifs (& = 0.05), une comparaison des 
moyennes a l’aide du test Tukey HSD a été effectuée 
entre les populations. 

Les données des trois populations ont ensuite été 
analysées pour vérifier l’existence d’une relation entre 
chaque variable et l’éloignement par rapport a la ligne 
d’eau. La meilleure régression, linéaire ou quadratique, 
a été sélectionnée a chaque population et pour chaque 
variable afin d’expliquer la relation. Les critéres de 
sélection pour la meilleure régression était la puissance 
du coefficient de régression, le raisonnement de |’ inter- 
prétation mathématique et la signification biologique. 


Résultats 
Mise a jour des données sur les effectifs, les localisa- 
tions et les superficies des populations 

Laster était présent a toutes les stations revisitées 
sauf deux. De plus, deux stations n’ont pas été revis- 
itées di aux contraintes de temps (Pointe de I’Est) et 
a l’établissement d’une résidence privée (Bassin aux 
huitres — secteur centre). Le tableau 1 présente les pop- 
ulations et leurs effectifs en comparaison des occur- 
rences notées dans Duclos (2004), Houle et al. (2002) 
et Gilbert et al. (1999). Les études de Houle et Haber 
(1990) et Gagnon et al. (1995a, 1995b) sont incluses 
dans Gilbert et al. (1999). 

Premiérement, il est 4 noter que |’étude de Houle et 
al. (2002) divise les populations (25 stations) alors que 
Gilbert et al. (1999) tend a les regrouper (8 occur- 
rences). Duclos (2004) a utilisé la numérotation de 
Houle et al. (2002). La présente étude propose une nou- 
velle définition des populations issues des stations de 


2005 


Houle et al. (2002). Quelques stations qui n’ étaient pas 
clairement divisées sur le terrain ont été regroupées. 
Ainsi, |’étude actuelle définit 18 populations nommeées 
selon des références géographiques plutot que num- 
érotées arbitrairement. Les informations relatives a 
chaque populations sont présentées ci-apres: 


Cap de l’hépital 
Classe 4 (+ 1500 individus) 
Localisation : N 47°25.112' et N 47°25.100' 

W 61°53.808' W 61°53.933' 
Cette population a décliné depuis les premiéres observations 
de Houle et al. (2002). Un ensablement du site a entrainé une 
diminution considérable de |’effectif en 2001. Les observa- 
tions récentes (Duclos 2004 et cette étude) montrent que la 
population a survécu a cette perturbation, mais que son 
effectif n’est pas encore retourné aux valeurs déja observées. 


Barachois, Chemin Poirier 
Classe | (4 individus) 
Localisation : N 47°25.180' 

W 61°9 1-971" 
Petite population spatialement distincte de la grande popula- 
tion Barachois. Seulement quelques individus répartis trés 
localement. Houle et al. (2002) ont observé une population de 
classe 3 en 1999 et 2000, alors qu’en 2001 la population était 
de classe 2. Ce présent relevé note une population de classe 1. 
Ainsi, cette population montre une tendance continuelle au 
déclin depuis les premiéres observations. 


Barachois 
Classe 5 (+ 63 000 individus) 
Localisation : N 47°25.440' a N 47°25.365' 

W 61°52.015' W .61°51.689' 
La population commence au bout d’un chemin (début de la 
grande plage) et est présente dans les premiers metres de 
végétation a l’interface entre la grande plage et le barachois. 
La population démontre une grande variabilité phénotypique. 
A la pointe du barachois les individus sont plus hauts et rami- 
fiés et sont répartis en tiges isolées. Vers le début de la grande 
plage, les individus sont petits, peu ramifiés et regroupés en 
touffes denses. L’ habitat est soumis a des perturbations natu- 
relles (inondation, ensablement) et anthropiques (VTT — 
véhicules tout-terrain). En comparaison avec les études pré- 
cédentes, l’ordre de grandeur de cette importante population 
apparait relativement stable. Gilbert et al. (1999) décrivait une 
population de classe 5, alors que Houle et al. (2002) et Duc- 
los (2004) l’ont classée 6. Dans la présente étude, la popula- 
tion est reclassée 5. Cette population a peut-étre été sous- 
évaluée ou surévaluée en raison de la difficulté d’ estimation 
due a la superficie et l’effectif considérables. Ainsi, en |’ ab- 
sence d’une tendance continue entre les études, il est diffi- 
cile de poser des conclusions quant a la dynamique de cette 
population. Le protocole de suivi a long terme inclura ce site 
afin de déterminer si les différences observées proviennent 
de différentes méthodes d’ estimation ou de tendances réelles. 


Etang du Nord 

Classe 0, aucun individu retrouvé a ce site (station 13 de 
Houle et al. (2002)). En 2001, Houle et al. (2002) ont constaté 
un déclin de cette population qui passait de la classe 3 a la 
classe 1. Trois ans plus tard, il est possible que cette population 
ait décliné jusqu’a |’ extinction. 


DE LAFONTAINE: L’? ASTER DU SAINT LAURENT AUX ILES-DE-LA-MADELEINE 


Pee 


Havre aux Basques 
Classe 5 (+ 64 000 individus) 
Localisation : N 47°18.394' a N47°17.465' 

W 61°56.329' W61°56.080' 
La population est répartie en deux zones le long de la lagune. 
Dans la premiére zone, quelques tiges isolées sont retrouvées 
dans les sites favorables. Le littoral est tranché par I’ érosion 
et les dép6ts de zostére a plusieurs endroits ce qui diminue le 
nombre de sites favorables a l’établissement de l’aster. La 
seconde zone compte une plus grande densité d’individus et 
est caractérisée par des tiges plus ramifiées et plus hautes. Le 
milieu est apparu moins perturbé. Ici encore, les estimés ne 
sont pas constants entre les études et aucune tendance ne peut 
étre décelée. Comme la population du Barachois, celle-ci sera 
inclue dans le suivi. 


Pointe des véliplanchistes 
Classe 3 (+ 200 individus) 
Localisation : N47°16.928' a N47°15.816' 
W61°55.808' W61°55.632' 
La population peu nombreuse couvre une vaste étendue tres 
perturbée par les inondations. Par rapport a la seule autre étude 
qui traite spécifiquement de cette population (Houle et al. 
2002), Veffectif a diminué considérablement passant d’une 
population de classe 6 a une population de classe 3. Ceci 
indique un changement certain de l’effectif de cette popula- 
tion indépendant des méthodes utilisées pour |’estimation. 
Cependant, puisque aucune autre étude ne s’attarde spéci- 
fiquement a cette population, on ne peut déterminer s’il s’agit 
dune véritable tendance au déclin. 


Pointe aux canots 
Classe 3 (+ 100 individus) 
Localisation : N47°15.959' a N47°16.016' 
W61°58.569' W61°58.694' 
La petite population est soumise a des perturbations (inonda- 
tion, dépdt de zostére). Lors d’une comparaison entre les sites 
de Pointe aux canots et Havre aux Basques, Reynolds et Houle 
(2002) observaient que la population de Pointe aux canots 
était dans une baie protégée. Cependant, lors du relevé de 
2004, j'ai pu observer que de nombreux individus étaient 
ensevelis sous d’épais dépdts de zostére. L’essentiel de la végé- 
tation croissait immédiatement apres la ligne d’eau sur le lit- 
toral, ne laissant que peu de micro-sites favorables a |’ étab- 
lissement de I’aster. L’étude de Houle et al. (2002) comptait 
beaucoup plus d’individus que |’ étude de Duclos (2004) et la 
présente étude. En effet, selon Houle et al. (2002), la popula- 
tion était de classe 6 alors que les relevés suivants présentent 
une population de classe 3. Tel que mentionné pour la popula- 
tion précédente, ceci indique un changement certain de |’ ef- 
fectif indépendant des méthodes utilisées pour I’ estimation. 
De plus, puisque la diminution est constante au cours de deux 
études subséquentes, il semble que la population soit en déclin. 


Dune de l’ouest 

Classe 3 (+ 100 individus) 

Localisation : N47°19.429' a N47°19.377' 
W61°57.448' a W61°57.482' 

L’étude de Houle et al. (2002) recensait plus de 2 millions 

d’individus dans cette population et l’occurrence polygo- 

nale notée par le centre de données sur le patrimoine naturel 

du Québec (CDPNQ) (Gagnon et al. 1995b) était de superficie 

considérable. Or, en 2004, seulement une centaine d’ individus 


560 


ont été retrouvés regroupés dans une zone trés restreinte. Le 
milieu ne présente pas une inclinaison importante et il s’agit 
plutot d’une vaste plaine hautement productive. Lors de la 
visite, toute cette plaine était inondée et des espéces parfois 
retrouvées en compétition avec |’aster et non aquatiques (e.g., 
potentille ansérine, Potentilla anserina L.) étaient submergées. 
Par conséquent, l’habitat propice a la croissance de I’aster 
n était que peu présent sur le site. Il est encore une fois diffi- 
cile d’ observer une tendance continue puisque seule |’ étude de 
Houle et al. (2002) sert de point de comparaison pour cette 
population. Cependant, une diminution allant de plus de deux 
millions 4 seulement une centaine d’individus est jugée sub- 
stantielle et méme dramatique. Un autre relevé devrait étre 
effectué dans les prochaines années afin de suivre I’état de 
cette population. 


Etang des caps 

Classe 6 (+ 240 000 individus) 

Localisation : N47°16. 598' a N47°16.690' 
W61°59.155' W61°58.416' 

Lessentiel de cette population est situé sur un ilot dans la 

lagune du Havre aux Basques accessible a gué. Depuis I’ étude 

de Houle et al. (2002), la population est caractérisée par un 

effectif tres élevé. De toutes les populations visitées, celle-ci 

est présente |’ effectif le plus important. 


Baie du portage 
Classe | (8 individus) 
Localisation : N47°15.381' 

W61°56.423' 
Cette population n’avait pas été revisitée depuis |’étude de 
Gagnon et al. (1995b) ou l’effectif exact n’ était pas indiqué. 
Le relevé de 2004 constitue donc la premiére évaluation de 
leffectif de cette petite population. 


Baie de Grosse-lle 

Classe 5 

Localisation : N47°37.366' a N47°37.428' 
W61°32.401' a W61°32.434' 

La population est située le long d’un sentier de VTT. Le gradi- 

ent de perturbation / compétition est donc entretenu par le pas- 

sage récurrent de VTT. Leffectif de cette population a aug- 

menté depuis la derniére observation par Houle et al. (2002). 

I] est possible que la popularité des VTT sur Grosse-Ile (Alain 

Richard. Attention Frag’ Iles, communication personnelle) en- 

courage la croissance démographique de la population d’ aster 

par le passage de nombreux appareils. 


Baie Clarke 

Classe 4 (+ 1000 individus) 

Localisation : N47°37.394' a N47°37.358' 
W61°28.331' W61°28.368' 

Cette population est également située le long d’un sentier de 

VTT et les sites favorables sont entretenus par leur passage 

récurrent. La population est stable depuis I’ étude de Houle et 

al. (2002) et l’effectif a augmenté depuis les premiéres obser- 

vations en 1995 (Gilbert et al. 1999). 


Pointe Old-Harry 

Classe 0. Bien que I’habitat semble propice, aucun individu 
n’a été retrouvé a ce site (station 6 de Houle et al. (2002)). Le 
suivi de Houle et al. (2002) notait une population stable de 
classe 2. Aprés une recherche exhaustive autour de la coordon- 
née géoréférencée fournie dans Houle et al. (2002), aucun 
individu n’a été retrouvé. Houle et al. (2002) notaient que 
ensemble de la végétation était fauchée pour entretenir un 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


espace de stationnement, il est possible que la taille continuelle 
de cette plante annuelle a empéché |’ établissement du stade 
adulte reproducteur menant ainsi cette petite population a 
extinction. 


Bassin aux Huitres, bassin est 
Classe 3 (+ 500 individus) 
Localisation : N47°33.543' 

W61°30.304' 
La population se trouve dans le gravier du stationnement. 
Houle et al. (2002) avait noté la présence de nombreuses tiges 
d’aster, constituant l’essentiel de la population prés d’un 
étang. Aujourd’ hui, l’étang s’est eutrophié et les algues vertes 
ont envahi les sites favorables a l’aster. La tendance 4 I’ eu- 
trophisation avait débuté lors du dernier relevé de Houle et al. 
(2002) (Gilles Houle, communication personnelle). Dans la 
présente étude, aucun aster n’a été retrouvé pres de 1|’étang. 
Duclos (2004) avait concentré l’effort d’échantillonnage prés 
de cet étang et n’avait pas noté la présence de I|’aster dans le 
gravier (Genevieve Duclos, communication personnelle). La 
perte de cette partie de la population a entrainé un déclin, 
l’ensemble de la population passant de la classe 6 (Houle et al. 
2002) a la classe 3 en 2004. 


Bassin aux Huitres, club vacance 
Classe 2 (+ 50 individus) 
Localisation : N47°33.366' 

W61°30.936' 
Les individus de cette population croissent a travers les galets 
d’une mise a |’eau pour embarcations nautiques. La popula- 
tion a conservé I’ effectif noté par Houle et al. (2002). 


Bassin aux Huitres, centre 

La population n’a pas été revisitée, une habitation privée avait 
été batie a proximité de la station de Houle et al. (2002). 
Cependant, lors du dernier relevé de Houle et al. (2002), il ne 
restait qu’un seul individu et Duclos (2004) n’en a noté aucun. 
Il est donc trés possible que cette population soit maintenant 
éteinte. 


Bassin aux Huitres, bassin ouest 
Classe 3 (+ 500 individus) 
Localisation : N47°32.613' a N47°32.656' 
W61°31.848' W61°31.946' 
Les individus sont répartis le long d’une bande a I’ interface 
entre un milieu dunaire et une zone fréquemment inondée. La 
population semble en déclin continuel. Gilbert et al. (1999) et 
Houle et al. (2002) notaient une population de classe 6; dans 
Duclos (2004) elle passait vers la classe 4 et dans la présente 
étude une population de classe 3 est observée. Houle et al. 
(2002) ont noté une forte érosion de |’ habitat de l’aster et une 
diminution importante en 2001 ne laissant qu’ une population 
de classe 2. Il est donc possible que cette population soit 
soumise a de fortes perturbations cycliques et que l’effectif de 
cette population soit tres variable. 


Pointe de lest 

Cette population n’a pas été revisitée 

Elaboration d’un protocole de suivi simple 

Les trois populations retenues pour |’élaboration du 
protocole de suivi sont Barachois (BAR), Havre aux 
Basques (HAB) et Etang des caps (EDC). Les localisa- 
tions exactes de chaque transect permanent ont été 
identifiées a l'aide d’un GPS et sont présentées au 
tableau 2. 


2005 DE LAFONTAINE: L? ASTER DU SAINT LAURENT AUX ILES-DE-LA-MADELEINE 56] 


TABLEAU 2. Localisations des 15 transects permanents installés aux trois populations sélectionnées pour le suivi a long 


terme. 

Population Transect Latitude Longitude 

Barachois (BAR) BARI N 47°25.369' W 61°51.682' 
BAR2 N 47°25.400' W 61°51.703' 
BAR3 N 47°25.432' W 61°S 1.763" 
BAR4 N 47°25.448' W 61°51.962' 
BARS N 47°25.442' W 61°52.012' 

Havre aux Basques (HAB) HAB1 N 47°18.388' W 61°56:332' 
HAB2 N 47°18.245' W 61°56.290' 
HAB3 N47 790 W 61°56.212' 
HAB4 N 477732 W 61°56.250' 
HABS N 47°17.686' W 61°56.241' 

Etang des caps (EDC) EDC] N 47°16.707' W 61°58.905' 
EDC2 N 47°16.719' W 61°58.877' 
EDG3 N 47°16.717' W 61°58.871' 
EDC4 N 47°16.702' W 61°58.764' 
EDGS N 47°16.687' W 61°58.626' 


Description de la situation actuelle aux populations 
sélectionnées pour le suivi 

Des différences significatives entre les populations 
ont été notées pour toutes les variables a I’ étude (Tableau 
3). Aussi, pour quatre des: six variables étudiées, il 
existe des différences entre les cinq transects d’une 
méme population. L’absence de différence significative 
pour les deux autres variables serait attribuable au 
manque de degrés de liberté associé au modéle con- 
tenant de nombreuses données manquantes. En effet, 11 
n’y avait aucune valeur de hauteur ou de nombre de 
capitules associés aux quadrats ou il n’y avait pas 
d’aster. La figure 1A montre que la population Etang 
des caps (EDC) est la plus dense (40.00 + 6.23 tiges 
d’aster / 0.5 m7’) et est significativement supérieure a 
celle de Havre aux Basques (HAB) (8.90 + 2.00 tiges 
d’aster / 0.5 m’). La population Barachois (BAR) n’est 
pas différente des deux autres avec une valeur inter- 
médiaire (21.90 + 10.51 tiges d’aster / 0.5 m7’). La fig- 
ure 1B montre que la population Etang des caps a un 


pourcentage de couverture par I’aster significativement 
supérieur (6.00 + 0.98 %) par rapport aux deux autres 
sites (HAB : 2.60 + 0.62 % et BAR : 1.33 + 0.40 %). 
Cependant, a la figure 1c, on constate que la population 
Etang des caps a un pourcentage de couverture par 
l'ensemble de la végétation inférieur (16.77 + 2.75 %) 
par rapport aux deux autres populations (HAB : 52.57 
+ 5.96 % et BAR : 64.57 + 6.70 %). La hauteur des tiges 
d’aster ainsi que le nombre de capitules par individu 
présentent le méme patron de variation entre les trois 
sites (Figures | D et E) : Havre aux Basques a les valeurs 
les plus élevées (respectivement 18.86 + 1.08 cm et 
48.50 + 8.10 capitules par plant). Etang des caps a des 
valeurs intermédiaires significativement différentes des 
deux autres populations (respectivement 11.63 + 0.74 cm 
et 25.93 + 3.17 capitules par plant). Enfin, Barachois 
a les valeurs les plus basses (respectivement 8.41 + 
0.84 cm et 6.12 + 1.41 capitules par plant). Enfin, la 
figure IF montre que la richesse spécifique est plus 
élevée a Havre aux Basques (7.17 + 0.59 especes / 


TABLEAU 3. Résultats de | ANOVA nichée pour évaluer les différences inter-populations et inter-transects pour les variables. 


Les valeurs P en caractéres gras sont significatives. 


\Variables Facteurs Valeurs F Valeurs P 
Nombre de tiges d’ aster Population 6.098 0.0035 
Transect (population) 3.039 0.0016 
Pourcentage de couverture par |’ aster Population 15.069 <0.0001 
Transect (population) 227 0.0009 
Pourcentage de couverture par la végétation Population 42.529 <0.0001 
Transect (population) 8.377 <0.0001 
Hauteur des tiges d’aster Population 24.925 <0.0001 
Transect (population) 1.831 0.066 
Nombre de capitules par aster Population 9.612 0.0003 
Transect (population) 0.878 0.574 
Richesse spécifique Population 3.208 0.046 
Transect (population) 7.441 <0.0001 


562 THE CANADIAN FIELD-NATURALIST Vol. 119 


as N (ee) aS Oo 
(a) oO (s) (2) (jo) 


Nombre de tiges d'aster / 0.5 m2 


(2) 


BAR HAB EDC 


Pourcentage de couverture 
par la végétation 


BAR HAB EDC 


oS 
oO 
eh 


No 
(=) 
it 


———— 


=< 
i=) 


Nombre de capitules par plante 
w 
oO 


j=) 


BAR HAB EDC 


Populations 


Pourcentage de couverture par l'aster 


BAR HAB EDC 


25 


Hauteur des tiges d'aster (cm) 


BAR HAB EDC 


Richesse spécifique 
(Nombre d'espéces / 0.5 m2) 


OF NY WF ATA DN CO CO 


BAR HAB EDC 


Populations 


FicureE |. Variables a 1’ étude sur chacun des sites (moyenne par quadrat de 0.5 m? + SE). Les lettres différentes au-dessus 
des colonnes représentent des valeurs moyennes significativement différentes (P < 0.05) entre les sites. 


0.5 m?) qu’a Etang des caps (5.77 + 0.47 espaces / 
0.5 m’), alors qu’elle est intermédiaire 4 la popula- 
tion Barachois (6.53 + 0.54 espéces / 0.5 m7). 


Quoique 1’ ANOVA nichée démontre qu’il existe 


des différences significatives entre les transects a 
chacun des sites et que ces transects ne peuvent donc 
€tre considérés comme de véritables réplicats, on 
observe des relations significatives entre les variables 
et la distance par rapport a la ligne du rivage pour les 
populations HAB et EDC (Figure 2). La population 


Barachois ne présente aucune tendance en fonction 
de la distance par rapport a la ligne du rivage. Les 
résultats complets sont présentés au tableau 4 et 
seules les régressions sélectionnées sont présentées 
dans le texte. Les figures 2A et B montrent que le nom- 
bre d’aster et le pourcentage de couverture d’aster 
varient de fagon quadratique (respectivement, HAB 
R? = 0,295, P= 0.09 6 EDC R? = 0.218, P= 0036 
HAB.R? = 0.292, P= ,0.0095.ct EDC R= 0 2e0, 
P =0.012) avec un mode aux distances intermédiaires 


2005 


DE LAFONTAINE: L’ ASTER DU SAINT LAURENT AUX ILES-DE-LA-MADELEINE 


Pourcentage de couverture 
par la végétation 


[o)) 
S 
O 


Nombre de capitules par plante 
NO 
(a) 

Q 
\ 
\ 
4 
| 
| 
| 
“a 


Richesse spécifique 
(nombre d'espéces / 0.5m2) 


0,0 0,5 1,0 1,5 2,0 2,9 3,0 


563 


= 
oO 
se 
e) 


S 
\ 
\ 


Oo 
aes 


Hauteur des tiges d'aster (cm 
% 
d 


i=) 
uc 
4 
+ 
71 


[oe] 


(o>) 
re 


Ba 
4 


N 
1 


Distance par rapport a la ligne du rivage (m) 


—@® BAR 
O-- HAB 
—w— EDC 


FIGURE 2. Variation des variables a |’ étude le long du gradient topographique a chaque site. BAR’, HAB* ou EDC’ indique une 
régression significative , suivie de L lorsque linéaire ou Q lorsque quadratique. 


par rapport a la ligne du rivage (i.e. ces deux vari- 
ables augmentent en s’éloignant du rivage pendant 
1.5 m puis diminuent). Par contre, a la figure 2c, il 
est démontré que le pourcentage de couverture de 
l’ensemble de la végétation augmente de facon liné- 
aire et positive en s’éloignant de la ligne du rivage 
(ABR? = 0.247, P:=/0:0052 et BDC R? = 0177, 
P =0.021). A la figure 2d, on voit que la hauteur des 
tiges d’aster varie de facon quadratique (HAB 
R? = 0358,,P =0.015,et EDC R2 = 0.268. P’= 0.015): 
La figure 2e montre que le nombre de capitules varie 


aussi de fagon quadratique A la population Etang des 
caps (R? = 0.211, P = 0.041), alors qu’a la figure 2f il 
est montré que la richesse varie de fagon quadratique 
a Havre aux Basques (R? = 0.232, P = 0.028). 


Discussion 
Mise a jour des données sur les effectifs, les localisa- 
tions et les superficies des populations 

L’étude de Gilbert et al. (1999) regroupe toutes les 
populations entourant la lagune du Havre aux Basques 
(Havre aux Basques, Pointe des véliplanchistes, Pointe 


564 THE CANADIAN FIELD-NATURALIST 


Vol. 119 


TABLEAU 4. Régressions (linéaires : y = 6,x + ,; et quadratiques y = f,x° + 6.x + f,) des différentes variables (y) en fonction 
de la distance par rapport a la ligne du rivage (x). Les valeurs en caractéres gras sont significatives (P < 0.05) et * indique 


que la régression a été sélectionnée lorsque les régressions linéaires et quadratiques étaient significatives. 


Variables Populations 


Nombre de tiges d’ aster BAR 
HAB 
EDC 
Pourcentage de couverture par |’ aster BAR 
HAB 
EDC 
Pourcentage de couverture par la végétation BAR 
HAB 
EDC 
Hauteur de I’ aster BAR 
HAB 
EDC 
Nombre de capitules par plant BAR 
HAB 
EDC 
Richesse spécifique BAR 
HAB 


BDC 


aux canots, Dune de l’ouest, Etang des caps et Baie du 
portage). Or, une seule station de Houle et al. (2002; 
station 19) montre un effectif dix fois supérieur a celui 
observé pour |’ensemble des populations entourant la 
lagune regroupées dans Gilbert et al. (1999). II est pos- 
sible que Houle et al. aient surestimé les populations 
par rapport aux autres études puisque dans |’ étude de 
Houle et al. (2002), tous les relevés ont été effectués 
alors que les individus étaient au stade végétatif. Par 
conséquent, il est possible que Houle et al. aient inclus 
des plantules n’ayant pas encore émergé. Toutes les 


autres études se déroulant au mois de septembre, pen-’ 


dant la floraison, rapportent des effectifs un peu plus 
faibles que ceux de Houle et al. (2002). Cependant, il 
parait peu probable que cette différence dans la phé- 
nologie explique a elle seule une diminution aussi 
importante que celle observée, par exemple, a la pop- 
ulation de la dune de l’ouest (de plus de 2 millions a 


Régression Coefficients de régression (RC) Valeurs P 
Linéaire 0.0097 0.605 
Quadratique 0.0366 0.605 
Linéaire 0.0063 0.676 
Quadratique 0.295* 0.009* 
Linéaire 0.0006 0.894 
Quadratique 0.218* 0.036* 
Linéaire 0.0029 O7TT 
Quadratique 0.0300 0.663 
Linéaire 0.0127 0.553 
Quadratique 0.292* 0.0095* 
Linéaire 0.045 O57 
Quadratique 0.280* 0.012* 
Linéaire 0.093 0.102 
Quadratique 0.101 0.238 
Linéaire 0.247* 0.0052* 
Quadratique 0.249 0.021 
Linéaire 0.177% 0.021* 
Quadratique 0.179 0.070 
Linéaire 0.0069 Wig fey! 
Quadratique 0.0066 0.619 
Linéaire 0.221 0.027 
Quadratique 0.358* 0.015* 
Linéaire 0.110 0.074 
Quadratique 0.268* 0.015* 
Linéaire 0.0071 0.748 
Quadratique 0.134 0.365 
Linéaire 0.131 0.098 
Quadratique 0.194 0.129 
Linéaire 0.159 0.029 
Quadratique 0.211* 0.041* 
Linéaire 0.04 0.290 
Quadratique 0.065 0.404 
Linéaire 0.104 0.083 
Quadratique 0.232* 0.028* 
Linéaire 0.0092 0.613 
Quadratique 0.142 0.126 


une centaine d’individus). Dans un souci d’ uniformiser 
les futurs inventaires, les relevés subséquents devraient 
étre effectués apres la deuxieme semaine d’aoiat et avant 
la troisieme semaine de septembre afin de tenir compte 
uniquement des individus adultes ayant émergé. 

Les études de Gagnon et al. (1995a, 1995b) citées 
dans Gilbert et al. (1999) ont été réalisées au cours 
d’une année dont la période de croissance (mai a sep- 
tembre) était particulierement seche (Archives Nat- 
ionales d’information et de données climatologiques 
(Environnement Canada) données utilisées aux stations 
de Gaspé et Charlottetown (IPE) disponibles sur www. 
climate.weatheroffice.ec.gc.ca; tableau 5). Une étude 
sur les effets du stress hydrique sur différents stades de 
croissance de I’aster a démontré qu’une sécheresse avait 
un effet négatif sur l’établissement des plantules (Houle 
et Belleau 2000). Par conséquent, il est possible que les 
études de Gagnon et al. (1995a, 1995b) aient eu lieu 


2005 


DE LAFONTAINE: L’ ASTER DU SAINT LAURENT AUX ILES-DE-LA-MADELEINE 


565 


TABLEAU 5. Tableau des données de pluviométrie mensuelle pour les saisons de croissance des années 1995 et 2004 et les 
moyennes mensuelles aux stations météorologiques de Gaspé et Charlottetown (IPE). Source : Archives Nationales d’information 
et de données climatologiques (Environnement Canada). ND indique que les données ne sont pas disponibles. 


Site Période 
mai juin 
Gaspé 1995 45.0 S2e0) 
2004 86.0 67.6 
Moyennes 90.8 73.4 
Charlottetown 
(IPE) 1995 69.7 96.2 
2004 ND ND 
Moyennes 97.7 93:2 


pendant une année ou peu d’individus adultes avaient 
émergé entrainant des effectifs plus bas que la moy- 
enne. ; 

Au contraire, la présente étude s’est déroulée a la fin 
dune saison de croissance ou les précipitations ont été 
plus abondantes que la moyenne (Archives Nationales 
d’information et de données climatologiques (Envi- 
ronnement Canada) données utilisées a la station de 
Gaspé disponibles sur www.climate.weatheroffice.ec. 
gc.ca; tableau 5). Les niveaux d’eaux étant élevés, 
beaucoup de sites favorables a la croissance de |’aster 
étaient soit érodés (e.g., Pointe des véliplanchistes), 
soit perturbés par des dépéts de zosteére (e.g., Pointe 
aux canots) Ou soit completement inondés (e.g., Dune 
de l’ouest). Ceci pourrait avoir eu pour effet de dimi- 
nuer |’effectif des populations a des valeurs plus bass- 
es que la moyenne. 

Les VTT semblent avoir un impact positif sur la 
croissance démographique de deux populations (Baie 
de Grosse-lle et Baie Clarke). Ceci peut étre expliqué 
de deux fagons. Premiérement, le passage récurrent des 
VTT peut créer le gradient de perturbation physique 
nécessaire a limiter la croissance des compétiteurs de 
Paster du Saint-Laurent en permettant la croissance de 
celle-ci. La dynamique retrouvée le long des lagunes 
est conservée et les €vénements d’inondation sont rem- 
placés par le passage d’un VTT. Deuxiémement, les 
VTT circulant sur de longues distances et créant un 
mouvement de I’air par leur passage peuvent faciliter 
la dissémination des akhénes d’ aster qui pourra s’ éta- 
blir facilement sur le sol dénudé laissé par le passage 
des véhicules. 

En résumé, les populations Cap de l’hépital, Bara- 
chois, Havre aux Basques, Etang des caps, Baie de 
Grosse-lle, Baie Clarke et Bassin aux Huitres (club 
vacance) sont stables ou en croissance. Les populations 
Barachois (Chemin Poirier), Pointe des véliplanchistes, 
Pointe aux canots, Dune de l’ouest et Bassin aux Huitres 
(est et ouest) semblent décliner. Enfin, les populations 
Etang du Nord, Pointe Old-Harry, Bassin aux Huitres 
(centre) semblent éteintes. Par contre, puisqu’il de- 
meure difficile de départager entre une tendance con- 
tinue au déclin et une dynamique variable selon les 


Pluviométrie mensuelle (mm) 


juillet aout septembre 
82.0 Tp lee 28.6 
109.8 133.6 95:9 
LOT 7 912 70.0 
Se )y2 2953 62,5 
ND ND ND 
85.8 87.3 95.4 


conditions annuelles, un suivi des populations a long 
terme demeure I’ outil a privilégier. 


Elaboration d’un protocole de suivi simple 

Puisque plusieurs sites étaient fortement perturbés 
lors de la visite de 2004 et que plusieurs populations 
semblaient étre caractérisées par des effectifs plus 
faibles que ceux observés dans d’autres études (e.g., 
populations de Dune de l’ouest ou Pointe des véliplan- 
chistes), le suivi devra inclure une visite annuelle de 
toutes les populations. La revisite de l'ensemble des 
populations permettra de continuellement mettre a jour 
les données d’effectif, de localisation et de superficie 
des populations. Une revisite de ces populations pourra 
éventuellement permettre |’ établissement de nouveaux 
transects permanents sur des populations importantes 
mais ou il n’y avait pas de sites favorables en 2004. 
De plus, une attention particuliére pourrait étre portée 
sur les populations de Baie de Grosse-lle et de Baie 
Clarke qui subsistent et dont l’effectif semble méme 
augmenter grace a des perturbations d’origine anthro- 
pique liées au passage récurrent de VTT. Aucun tran- 
sect permanent n’a été établi sur ces sites puisque les 
piquets auraient certainement nuit au passage des VTT 
et auraient été immédiatement retirés. Cependant, cette 
dynamique liée a la perturbation d’origine anthropique 
est intéressante et mérite que l’on s’y attarde a I’ avenir. 

Enfin, on peut s’attendre 4 ce que la ligne du rivage 
soit dépendante de la quantité de précipitations recues. 
Par conséquent, il est fort possible que cette ligne 
varie et qu’il soit nécessaire de réajuster |’emplace- 
ment des transects au cours des prochaines années 
afin de couvrir adéquatement |’ habitat de l’aster. Une 
facgon de le faire est de visualiser chaque transect per- 
manent comme deux lignes paralléles séparées par 
1m et d’une longueur indéfinie dont le premier 
quadrat (de | m Xx 0.5 m) est a proximité de la ligne 
du rivage et le dernier est situé 2.5 m plus loin pour 
donner un transect d’échantillonnage d’une longueur 
totale constante de 3 m. Si, au cours des prochaines 
années, il est noté que la ligne du rivage se déplace, 
les transects installés actuellement serviront de bornes 
indiquant les deux lignes paralléles; il ne suffira que 


566 


de tirer des lignes continues jusqu’a la ligne du rivage 
pour y installer le premier quadrat. Si une telle dé- 
marche doit étre utilisée, il faudrait inclure dans le suivi 
la mesure de la distance de déplacement du transect 
par rapport au transect permanent original. 


Description de la situation actuelle aux populations 
sélectionnées pour le suivi 

Les résultats ont montré que les trois populations 
choisies pour le suivi a long terme présentent des diffé- 
rences significatives pour toutes les variables incluses 
dans le suivi. La population Etang des caps présente 
la plus forte densité et les individus de cette population 
couvrent un plus grand pourcentage de sol. Ces résul- 
tats soutiennent les estimés de l’effectif ot Etang des 
caps était considérée comme la seule population de 
classe 6. Ceci peut étre di a une faible compétition 
interspécifique, puisque les résultats de l’analyse ont 
montré qu’il s’agit du site ou le pourcentage de |’en- 
semble de la végétation et la richesse spécifique sont 
les plus faibles. Ceci indique donc une différence au 
niveau de la dynamique de cette population par rap- 
port aux deux autres populations. Cette dynamique 
favorise la croissance de I’aster par rapport aux autres 
especes. 

Les individus de la population Havre aux Basques 
ont atteint en 2004 une taille plus haute et possedent 
davantage de capitules alors que les individus de la 
population Barachois étaient, au contraire, les plus 
petits et les moins ramifiés. La densité de la popula- 
tion du Barachois est deux fois plus élevée que celle 
observée au Havre aux Basques. Il est intéressant de 
constater que toutes les autres variables mesurées a 
ces deux sites ne présentent pas de différences signi- 
ficatives. Ces deux populations semblent donc utiliser 
deux stratégies morphologiques distinctes. La popu- 
lation du Barachois double la quantité de tiges mais 
réduit la taille et le nombre de capitules de celles-ci, 
alors que la population du Havre aux Basques, moins 
dense est composée d’individus plus hauts et ramifiés. 
Ces deux stratégies permettent d’obtenir le méme 
pourcentage de couverture par |’aster. I] semble que 
la compétition interspécifique ne soit pas reliée a l’u- 
tilisation de l’une ou |’ autre de ces stratégies puisque 
les deux populations sont situées sur des sites ayant 
une méme richesse spécifique et un méme pourcent- 
age de couverture de végétation. L’utilisation de stra- 
tégies distinctes doit donc étre reliée a une variable qui 
n’a pas été mesurée dans le suivi telle que la salinité 
du substrat qui inhibe la germination (Houle et al. 
2001) ou une période de sécheresse en début de saison 
de croissance qui retarde |’établissement des plan- 
tules (Houle et Belleau 2000). 

Il appert que les trois populations possedent des 
différences qui renseignent sur |’ importance de la den- 
sité, la morphologie et habitat. Par conséquent, il est 
fortement suggéré de conserver ces trois populations 
et ces six variables dans le suivi a long terme afin de 
vérifier dans quelle mesure ces populations conservent 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


les mémes caractéristiques au cours des années sub- 
séquentes ou si la dynamique est sujette a changer (e.g., 
en fonction de la quantité de précipitations). Aussi, 
tel que mentionné plus haut, l’inclusion de nouvelles 
populations est recommandeée, particuliérement celles 
dont les effectifs varient entre les études (e.g., Dune de 
l’ouest, Pointe des véliplanchistes, Pointe aux canots). 

Pour quatre des six variables, il existe des diffé- 
rences significatives entre les cinq transects de chaque 
population. Ceci suggére que les transects ne sont pas 
des réplicats et qu’ils doivent plutét étre considérés 
comme différents micro-habitats. Par conséquent, il 
est important de conserver les quinze transects dans 
le suivi a long terme. Retirer un transect constituerait 
une perte significative d’information. Les données 
recueillies au cours des prochaines années du suivi, 
permettront en plus de comparer la progression d’un 
méme transect au cours du temps. Ceci permettra de 
voir comment chaque micro-habitat se transforme. 

Méme si les transects ne sont pas de véritables répli- 
cats, il est possible de discerner des relations signi- 
ficatives des différentes variables en fonction de la 
distance par rapport a la ligne du rivage pour deux 
populations. Le plan d’échantillonnage de transects 
organisés perpendiculairement a la ligne de rivage sur 
une distance de 3 m permet de couvrir le gradient de 
perturbation / compétition attendu sur deux sites. En 
effet, aux populations de HAB et EDC, le nombre de 
tiges et le pourcentage de couverture par l’aster de 
méme que les valeurs associées aux variables morpho- 
logiques (la hauteur des tiges et le nombre de capitules 
par individu) sont faibles pres de la ligne du rivage ou 
le stress lié a la salinité (Houle et al. 2001; Reynolds 
et al. 2001) et les perturbations (Reynolds et Houle 
2002) empéchent la croissance de |’aster. En s’éloig- 
nant du rivage, les conditions du milieu favorisent la 
croissance de |’aster mais demeurent trop restrictives 
pour la croissance d’espéces compétitrices. Le mode 
de la distribution se situe dans cette bande intermédi- 
aire. A une distance plus éloignée de la rive, les condi- 
tions deviennent favorables pour des espeéces plus com- 
pétitrices qui limitent la croissance de l’aster (Brumbt 
2001; Houle et Valéry 2003). Ce gradient est confirmé 
par le pourcentage de couverture par l’ensemble de la 
végétation qui augmente de fagon linéaire a partir de 
la ligne du rivage. La richesse spécifique suit la distri- 
bution quadratique a la population Havre aux Basques. 
Il semble que cette relation soit un artéfact de la relation 
entre diversité et productivité fréquemment observée 
(Grime 1973; Rosenzweig 1992; Huston et deAngelis 
1994: de Lafontaine 2004). En effet, le maximum de 
richesse est observé a la distance correspondant a des 
valeurs intermédiaires de pourcentage de couverture 
de l’ensemble de la végétation. 

Par contre, a la population Barachois, le plan d’é- 
chantillonnage ne permet pas de déceler le gradient 
de perturbation / compétition attendu. Le site est par- 
ticulier et les transects T4 et TS ont été positionnés un 


2005 


peu en retrait par rapport a la ligne du rivage puisque 
la plupart des individus ne sont pas situés dans I’ habi- 
tat typique de la bande de 3 m apres la ligne de rivage. 
Il est possible de redéfinir l’emplacement de ces tran- 
sects afin qu’ils correspondent plus adéquatement au 
gradient typique. Il se peut que cette démarche uni- 
formise les transects a ce site créant ainsi de véritables 
réplicats. Cependant, tant que ces transects seront peu- 
plés d’aster, il est plut6t conseillé de les conserver tels 
qu’ils sont présentement en tenant compte de cette 
particularité dans le suivi a long terme. 


Conclusions 

Cette étude définit 18 populations. Les 7 popula- 
tions Cap de l’hépital, Barachois, Havre aux Basques, 
Etang des caps, Baie de Grosse-lle, Baie Clarke et 
Bassin aux Huitres (club vacance) sont stables ou en 
croissance. Les 6 populations Barachois (Chemin 
Poirier), Pointe des véliplanchistes, Pointe aux canots, 
Dune de l’ouest et Bassin aux Huitres (est et ouest) 
semblent décliner. Enfin, les 3 populations Etang du 
Nord, Pointe Old-Harry, Bassin aux Huitres (centre) 
semblent éteintes ou tres pres de l’extinction. La popu- 
lation Pointe de l’est n’a pas été revisitée et cette étude 
présente le premier estimé:de la population Baie du 
portage. A partir des études qui ont été réalisées en 
utilisant différentes approches, 11 est difficile de dépar- 
tager entre de réelles tendances au déclin et une dy- 
namique en fonction des conditions changeantes du 
milieu. L’utilisation d’un suivi a long terme appliquant 
un protocole uniforme permettra de déterminer la pro- 
gression des différentes populations. 

Un protocole a été élaboré et le plan d’ échantillon- 
nage a été installé a trois populations : Barachois, 
Havre aux Basques et Etang des caps. Il est conseillé 
d’ installer le plan d’échantillonnage sur d’ autres sites 
importants ou les effectifs semblent en déclin mais 
sur lesquels 11 était impossible d’établir des transects 
en 2004 (Dune de l’ouest, Pointe aux canots, Pointe des 
véliplanchistes). Il serait également intéressant d’étu- 
dier la dynamique des populations d’aster perturbées 
par le passage récurrent des VTT (Baie de Grosse-Ile 
et Baie Clarke). Enfin, il est recommandé d’effectuer 
une visite annuelle des 18 populations. 

Au sein des trois populations sélectionnées pour le 
suivi, les résultats obtenus en 2004 montrent des dif- 
férences significatives pour toutes les variables consi- 
dérées. Aux trois sites, les transects présentent des diffé- 
rences entre eux et ne peuvent donc pas étre considérés 
comme des réplicats. A la lumiére de ces informations, 
il est recommandé de conserver les trois populations 
pour le suivi a long terme et de conserver les cing 
transects tels qu’ils se présentent actuellement. Le 
plan d’échantillonnage permet de déceler le gradient 
de perturbation / compétition attendu sauf a la popu- 
lation plus particuliére du Barachois. Ceci indique que 
pour les populations de Havre aux Basques et Etang 
des caps, ce plan d’échantillonnage permet de tenir 


DE LAFONTAINE: L? ASTER DU SAINT LAURENT AUX ILES-DE-LA-MADELEINE 


567 


compte de la variation par rapport a la ligne du rivage 
et devrait étre conservé comme tel. Cependant, les 
transects 4 et 5 de la population du Barachois ont été 
déplacés a cause de contraintes particuli¢res associées 
ace site. Ces transects devraient étre laissés ainsi et 
lanalyse devra en tenir compte. 


Remerciements 

Le soutien financier a été fourni par le ministére 
du développement durable, environment et parcs du 
Québec et Environnement Canada. Les données ont été 
fournies par le ministere du développement durable, 
environment et parcs du Québec . 


Littérature citée 

Brumbt, C. P. 2001. Effet de la compétition interspécifique 
sur l’aster du Saint-Laurent (Aster laurentianus Fernald) 
aux Iles-de-la-Madeleine, Québec. Mémoire de maitrise. 
Université Laval, Québec. 56 pages. 

COSEPAC. 2004. Espéces canadiennes en péril, mai 2004. 
Comité sur la situation des espéces en péril au Canada. 
57 pages. 

de Lafontaine, G. 2004. Richesse spécifique le long d’un gra- 
dient de production: utilisation-d’une approche multivariée. 
Mémoire de maitrise. Université Laval, Québec. 44 pages. 

Duclos, G. 2004. Suivi des populations de Symphyotrichum 
laurentianum, une plante endémique du sud du Golfe du 
Saint-Laurent. These d’ initiation a la recherche. Université 
de Moncton, Moncton. 35 pages. 

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Received 24 January 2005 
Accepted 13 October 2005 


Tree Recruitment Limitation by Introduced Snowshoe Hares, Lepus 
americanus, on Kent Island, New Brunswick 


TREVOR S. PETERSON, AKANE UESUGI, and JOHN LICHTER 


Biology Department and Environmental Studies Program, 6500 College Station, Bowdoin College, Brunswick, Maine 04011 
USA 


Peterson, Trevor, Akane Uesugi, and John Lichter. 2005. Tree recruitment limitation by introduced Snowshoe Hares, Lepus 
americanus, on Kent Island, New Brunswick. Canadian Field-Naturalist 1 19(4): 569-572. 


Species introductions often have negative consequences for native plant and animal communities of islands. Herbivores 
introduced to islands lacking predators can attain high population densities and alter native plant communities by selective 
consumption of palatable plants. We examined the legacy of the 1959 introduction of Snowshoe Hares (Lepus americanus) 
to Kent Island, New Brunswick, by reconstructing a history of tree recruitment on Kent Island and on nearby Outer Wood 
Island, which lacks Snowshoe Hares. Tree-ring records show pronounced recruitment peaks associated with farm abandon- 
ment in the 1930s for Kent Island and in the 1950s for Outer Wood Island. Following the introduction of Snowshoe Hares to 
Kent Island, tree recruitment plummeted and has remained low ever since. In contrast, trees continued to establish through- 
out the latter 20" century on Outer Wood Island. The high rates of seedling mortality on Kent Island associated with Snow- 
shoe Hare browsing coupled with high rates of canopy tree mortality threaten to degrade severely the forest of this important 
seabird nesting sanctuary. 


Key Words: Snowshoe Hare, Lepus americanus, invasive species, tree recruitment limitation, Kent Island seabird sanctuary, 


New Brunswick. 


Introduced herbivores often cause great damage to 
island ecosystems and potentially result in plant species 
extinction (Van Vuren and Coblentz 1987; Moran 1996; 
Coblentz 1990). The absence of significant predators 
allows introduced herbivores to reach high population 
densities and exert intense grazing pressure on poorly 
defended plants (Krebs et al. 2002; Donlan et al. 
2002). Because woody plants are most vulnerable to 
browsing during their early stages of development 
and growth (Clark et al. 1999; Liang and Seagle 2002), 
intense browsing may limit tree regeneration and allow 
less palatable species to proliferate (Rossow et al. 1997; 
Heinen and Currey 2000). Shifts in plant species dom- 
inance resulting from intense herbivory have been 
demonstrated in tropical, temperate, and boreal island 
ecosystems and indicate the community-level impacts 
of herbivore introductions or predator removal from 
islands (McLaren and Peterson 1994; Chouinard and 
Filion 2001; Terborgh et al. 2001). 

In 1959, Snowshoe Hares (Lepus americanus) col- 
onized Kent Island, an 80-hectare island in the Bay of 
Fundy, New Brunswick, following intentional introduc- 
tion to Hay Island, a nearby island connected to Kent 
Island at low tide (C. Huntington, personal communi- 
cation). Subsequently, estimates of the hare popula- 
tion in various years have ranged between 50 and 500 
individuals based on mark-recapture techniques (N. 
Wheelwright, Bowdoin College; personal communica- 
tion). Snowshoe Hares are generalist herbivores that 
feed on a variety of grasses and shrubs during the sum- 
mer, but on tree bark, foliage, and twigs during the win- 
ter (de Vos 1964). Diet selection studies have demon- 


strated that hares preferentially browse deciduous spec- 
ies, but may rely on conifers during winter or during 
periods of high population density if preferred forage 
is not available (Aldous and Aldous 1944; de Vos 1964; 
Parker 1984). Snowshoe Hares often avoid White Spruce 
(Picea glauca) when other browse is available, presum- 
ably because of high levels of camphor in the foliage 
(Sinclair et al. 1988; Rangen et al. 1994; Heinen and 
Currey 2000). The concentration of secondary com- 
pounds in vegetation may also influence diet selection 
(Fox and Bryant 1984; Rousi 1997; Rodgers and Sin- 
clair 1997). Paper Birch (Betula papyrifera) saplings, for 
example, can effectively deter Snowshoe Hare brows- 
ing with high concentrations of secondary compounds 
(Bryant 1981). 

Although Snowshoe Hare browsing of trees is gen- 
erally limited to heights less than | m, hares are impor- 
tant herbivores in boreal forests, and considerable dam- 
age to seedlings and saplings in forest plantations has 
been attributed to hares (Aldous and Aldous 1944; Holl 
and Quiros-Nietzen 1999; Rao et al. 2003). Based on 
Kent Island’s high population of Snowshoe Hares and 
on observations of intense browsing of tree seedlings, 
we hypothesized that browsing by hares effectively 
halted tree recruitment since their introduction in 1959. 
To document the impacts of hare browsing on Kent Is- 
land, we reconstructed a tree recruitment history based 
on tree-ring records for Kent Island and for nearby 
Outer Wood Island, which lacks Snowshoe Hares. We 
also compared current tree recruitment rates on the 
two islands by estimating seedling and sapling densi- 
ties in randomly located plots. 


569 


570 


Study Area 

Kent Island is at the southern end of the Grand 
Manan archipelago in the Bay of Fundy, New Bruns- 
wick (44°35'N, 66°46'W). The island is an important 
nesting area for Leach’s Storm Petrel (Oceanodroma 
leucorhoa) and Savannah Sparrow (Passerculus sand- 
wichensis). Since 1935, Bowdoin College of Bruns- 
wick, Maine, has operated a biological station on Kent 
Island. In 1959, Snowshoe Hares were intentionally 
introduced to nearby Hay Island, and soon thereafter 
crossed the tidally exposed land bridge to Kent Island. 
Hare populations have persisted on both islands since 
that time. Muskrats (Ondatra zibethicus) were intro- 
duced earlier in 1941 (Huntington 1956). The forest 
community contains Balsam Fir (Abies balsamea), 
White Spruce, Red Spruce (Picea rubens), Yellow Birch 
(Betula allegheniensis), Heart-leaved Birch (Betula 
cordifolia), and Mountain Ash (Sorbus americanus). 
A mixed forest dominated by White Spruce and Bal- 
sam Fir covers approximately two-thirds of the north- 
ern half of the island, whereas grassland covers the 
southern half of the island. In the early 20" century, 
much of the grassland was either cultivated or used for 
hayfield and pasture (Gleason 1937). Today seedlings 
and saplings of any tree species are rare on Kent Island. 
Those that are found show a pronounced browse line 
up to 60 cm in height. Canopy trees are suffering high 
rates of mortality for unknown reasons, and the gaps 
formed by tree death and windthrow are rapidly colo- 
nized by Red Raspberry (Rubus idaeus) and Wood 
Fern (Dryopteris spinulosa) rather than trees. 

Outer Wood Island (44°37'N, 66°49'W) is approxi- 
mately 6 km northwest of Kent Island. Forest covers 
its central area, which is surrounded by grasslands sim- 
ilar to Kent Island. White Spruce and Red Spruce are 
the dominant canopy species with Balsam Fir, Yellow 
Birch, and Mountain Ash important locally. Outer 
Wood Island lacks Snowshoe Hares, but, like Kent 
Island, has a large introduced Muskrat population. 
Outer Wood Island was used for sheep pasture through- 
out the early and middle 20" century (Ingersoll 1991). 


Methods 

In March 2002, an attempt to eradicate the Snow- 
shoe Hare population on Kent Island was initiated, 
which resulted in a reasonably accurate population 
estimate. Eradication efforts have continued up to the 
present. Prior to the attempted hare eradication, we 
counted hare pellets in 132 plots (1 m x | m) randomly 
located within forests on Kent Island to document 
hare distribution on the island, as described by Krebs 
et al. (2001). | 

To describe the composition of the forests, we exam- 
ined trees occurring along the length of nine randomly 
located 2 m x 50 m transects on each island. Canopy 
trees were identified to species and their diameter-at- 
breast-height (DBH) was recorded. To develop tree- 
recruitment histories, we collected increment cores 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


from White Spruce, Red Spruce, and Balsam Fir trees 
in nine randomly chosen circular plots on each island. 
Within each plot, the ten trees nearest the center and 
greater than S-cm DBH were cored at a height ranging 
between 30 and 50 cm from the ground. The species 
identity and DBH were recorded for all cored trees. 
We also took advantage of numerous treefalls that oc- 
curred during the winter of 2000-2001 by collecting 
basal discs from 33 trees. The discs and cores were 
sanded smooth, and the rings were counted using a 
binocular Leica dissecting microscope and a Velmex 
tree-ring measuring system. To account for early growth 
of the trees, we added five years to the tree-age esti- 
mates of basal discs and ten years to the estimates of 
increment cores (DesRochers and Gagnon 1997; Par- 
ent et al. 2000). 

On both islands, established seedlings and saplings 
were quantified in three 78.5-m7? (i.e., 5-m radius) sam- 
ple plots, located 25 m apart along each 50-m transect. 
Seedlings were defined as any stem less than | m in 
height and saplings were defined as stems greater than 
1 m in height and having a DBH less than 5 cm. Height 
measurements and extent-of-browsing estimates were 
recorded on isolated White Spruce seedlings on Kent 
Island and Outer Wood Island. 


Results 

The density of Snowshoe Hare pellets on Kent 
Island was 47.1 (4 6.2)/m? and ranged between 6 and 
110 pellets/m. Pellet densities were not significantly 
correlated with seedling abundances. No pellets were 
found on Outer Wood Island. After one month of hunt- 
ing in early 2002, 275 Snowshoe Hares had been killed 
on Kent Island, indicating a minimum density of 3.43 
hares per hectare. The hare population was not elimi- 
nated during this period of hunting, and the popula- 
tion has subsequently rebounded, although not to its 
maximum level. Eradication efforts are scheduled to 
continue in winter 2005-2006. 

The recruitment histories of canopy trees of the two 
islands show patterns of reforestation following farm 
abandonment and cessation of livestock grazing in the 
mid-20" century (Ingersoll 1991). Consistent with the 
timing of land-use change, peak tree recruitment oc- 
curred during the 1930s on Kent Island and during the 
1950s on Outer Wood Island (Figure 1). On Kent I[s- 
land, tree recruitment remained high through the 1940s 
but declined precipitously in the 1950s, whereas tree 
recruitment on Outer Wood Island was relatively high 
throughout the 1960s and 1970s, two decades after its 
peak (Figure 1). 

The abundances of saplings and established seed- 
lings (i.e., > 2 yrs old) reveal the disparity in recent 
tree recruitment on the two islands. Sapling abundance 
of all tree species was much greater on Outer Wood 
Island than on Kent Island (OWI = 29.4 + 13.3; 
KI =0.1 + 0:1; di =43; t=2.71; P =0,01) Gieure 2 
The abundance of established seedlings was similarly 


2005 


0.5 


PETERSON, UESUGI, and LICHTER: TREE RECRUITMENT LIMITATION 


571 


[1 Outer Wood Island 


0.4 2 Kent Island 


0.3 


0.2 


Proportion of tree 
establishment 


eae : ln lig oe | 


1860s 1870s 1880s 1890s 1900s 1910s 1920s 1930s 1940s 1950s 1960s 1970s 


Year 


FIGURE |. Tree recruitment history for Kent Island and Outer Wood Island, New Brunswick derived from tree-ring records. 
Data given as the proportion of the total number of canopy trees cored for each decade between 1860 and 1980. 
Snowshoe Hares were introduced to Kent Island in 1959. 


much greater on Outer Wood Island than on Kent Is- 
land (OWI = 14.1 + 3.5; KI = 0.3 + 0.2; df= 43; t=3.17; 
P =(0).003). All of the 58 spruce seedlings and saplings 
examined on Kent Island showed visible signs of 
browsing. Of these 58 trees, 40 were less than one 
meter in height, and 18 between | and 2 m tall. Of the 
40 less than | m in height, 24 were missing their api- 
cal meristem, whereas of the 18 ranging between | and 
2 m in height, only one had its apical meristem browsed. 
On Outer Wood Island, apical stems were present on 
all 60 of the seedlings sampled, and little or no evidence 
of browsing was present. 


Discussion 

The history of tree recruitment on Kent Island and 
Outer Wood Island demonstrates the long-term, neg- 
ative effects of intense Snowshoe Hare browsing on 
tree recruitment in an island ecosystem lacking mam- 
malian predators. The precipitous decline of tree re- 
cruitment on Kent Island during the 1950s suggests 
that the Snowshoe Hare population increased rapidly 
and that few established tree seedlings survived the 
onset of hare browsing. While we have no sequential 
estimates of the hare population density, the lack of 
saplings and established seedlings suggests that hares 
were numerous enough to limit tree recruitment severe- 
ly throughout the past forty-five years. Sustained brows- 
ing pressure can cause a shift toward less palatable 
species (McLaren and Peterson 1994; Heinen and Cur- 
rey 2000; Terborgh et al. 2001). Because all tree species 
on Kent Island are palatable to Snowshoe Hares, a 
shift to less palatable species would result in loss of 
forest. Currently as canopy trees die, Wood Fern and 
Red Raspberry replace them. A dense cover of Wood 


16 
(—~] Outer Wood Island 
12 5 Kent Island 
® 
2 8 
3 
4 
4 
0 


1-2 2-3 3-4 4-5 5-6 6-7 7-8 


Sapling Ht (m) 


FIGURE 2. Tree sapling abundance for Kent Island and Outer 
Wood Island, New Brunswick, given for height 
classes. 


Ferns probably also limits tree recruitment as Wood 
Ferns have been shown to limit the photosynthetic 
capabilities of coniferous seedlings (Starostina 1988). 

This study contributes to a growing body of research 
documenting pronounced ecosystem-level consequenc- 
es of herbivore introductions to islands (e.g., Van Vuren 
and Coblentz 1987; Donlan et al. 2003). Because Snow- 
shoe Hares are the only mammalian herbivores that 
browse trees on Kent Island, they are responsible for 
all of the browsing damage since their introduction. If 
the current hare eradication efforts are not successful, 
it is likely that this forest will become severely degrad- 
ed in the future as the canopy trees reach senescence. 
This outcome may negatively affect the suitability of 
the island as nesting habitat for Leach’s Storm Petrels, 


S12 


whose nest sites are currently concentrated in the forest- 
ed portions of the island. This would be unfortunate, 
because Kent Island provides vital and well-document- 
ed nesting habitat for Leach’s Storm Petrel (Gross 1935; 
Griffin 1940; Huntington 1956, 1959). Successful erad- 
ication of Snowshoe Hares from Kent Island would 
provide a valuable opportunity to document whether 
removal of the dominant herbivore will allow the for- 
est community of the island to recover from nearly 
half a century of suppressed tree recruitment. 


Acknowledgments 

We sincerely thank Nat Wheelwright, Barry Logan, 
and Chuck Huntington for thoughtful comments on 
this manuscript, and Sarah Rodgers, Jaret Reblin, and 
Peter Hill for help in the field. This is contribution 
number 154 from the Kent Island Scientific Station. 


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Received 11 January 2005 
Accepted 26 September 2005 


Travel Rates of Wolves, Canis lupus, in Relation to Ungulate Kill 
Sites in Westcentral Alberta 


GERALD W. Kuzyk!, CHRISTOPH ROHNER!, and FIONA K. A. SCHMIEGELOW! 


'Department of Renewable Resources, University of Alberta, Edmonton, Alberta T9H 4N1 Canada 
* Corresponding author: gkuzyk @ualberta.ca 


Kuzyk, Gerald W., Christoph Rohner, and Fiona K. A. Schmiegelow. 2005. Travel rates of Wolves, Canis lupus, in relation to 
ungulate kill sites in westcentral Alberta. Canadian Field-Naturalist 119(4): 573-577. 


Recent advancements in Global Positioning Systems (GPS) radiocollar technology permit analysis of fine-scale animal move- 
ments. We used concurrent aerial and GPS monitoring to determine winter travel rates of Wolves (Canis lupus) in relation to 
ungulate kill sites in managed forest landscapes in westcentral Alberta. Wolves preyed predominately on Moose (Alces alces) 
and travelled 4.2 times less when near ungulate kill sites than when away from them. As Wolves are thought to be an important 
factor in Woodland Caribou (Rangifer tarandus caribou) declines, information is needed to assess predation risk to Caribou 
from Wolves under a variety of landscape conditions. If Wolves have restricted movements near Moose kill sites, this may lead 
to decreased encounter rates with Caribou in systems where Moose are abundant. Deer (Odocoileus spp.) are probably an impor- 
tant component of this Wolf-prey system but little is currently known about this relationship. Projecting long-term implications 
of ongoing development activities requires a more detailed understanding of the responses of all species to landscape change. 


Key Words: Moose, Alces alces, Wolf, Canis lupus, predation risk, Woodland Caribou, Rangifer tarandus caribou, GPS data, Alberta. 


Wolves (Canis lupus) travel extensively to locate 
prey (Mech 1970). For example, when hunting Moose 
(Alces alces), Wolves may travel 30-50 kilometres per 
day (Mech 1966; Peterson 1977; Mech et al. 1998). 
Woodland Caribou (Rangifer tarandus caribou) are 
classed as threatened in Alberta (Edmonds 1998; Dzus 
2001) and Wolf predation is thought to be a major con- 
tributor to Caribou declines (Bergerud 1974; Edmonds 
1988; Hayes et al. 2003). It is hypothesized that Wood- 
land Caribou spatially separate themselves from Moose 
to avoid predation by Wolves (Bergerud and Elliot 
1986; Seip 1992). Wolves may alter their use of habitats 
in response to industrial development such as forest har- 
vesting (Kuzyk et al. 2004) and human infrastructure 
such as roads and trails can enhance Wolf movements 
(Formozov 1946; Thurber et al. 1994; Ciucci et al. 
2003). James (1999) reported that Wolves in winter trav- 
elled 2.8 times faster on a linear corridor than in the 
forest. Such enhanced mobility has implications for 
search efficiency and encounter rates of Wolves with 
prey species. Increased information on Wolf-prey sys- 
tems is essential for future Caribou conservation deci- 
sions (Hayes et al. 2003). Understanding Wolf travel 
rates in relation to ungulate kill sites offers one method 
of assessing predation risk to other prey species. 

When Wolves kill large prey such as Moose, they 
usually spend two to four days near the carcass (Peter- 
son 1977; Ballard et al. 1987; Mech et al. 1998; Hayes 
et al. 2000) whereas White-tailed Deer (Odocoileus 
virginianus) carcasses are generally handled in less 
than one day (Fuller 1989). After feeding on an ungu- 
late carcass, Wolves may travel several kilometres to 
rest in open sunny areas, where digestion may be opti- 
mized (Mech 1970). During one study, Wolves were 


found near Moose kills in 21 of 31 days of continuous 
monitoring (Mech 1966), but fine scale movements 
near the carcasses were not recorded. 

Observational studies of Wolves provide useful 
insight into Wolf hunting behaviour (Carbyn and 
Trottier 1988; Mech 1997) but there is little quantita- 
tive information on Wolf movements near kill sites, 
due to the technical difficulties of collecting such 
information (Mech 1995). Wolves have traditionally 
been studied using daily aircraft flights to relocate 
radiocollared Wolf packs, but this technique is limit- 
ed by daylight and favourable weather (Mech 1995). 

The abjective of this study was to combine Global 
Positioning Systems (GPS) radio-collar technology with 
concurrent aerial observations to provide detailed infor- 
mation on Wolf travel rates in relation to ungulate kill 
sites. Wolves feeding on Moose are predicted to restrict 
movements when near carcasses (Mech 1966; Mech 
1970) which should, theoretically, lessen predation risk 
to Caribou in systems where moose are abundant. Wolf 
packs feeding on deer should spend minimal time at 
kill sites (Fuller 1989), and more time travelling, which 
could result in increased predation risk to Caribou 
through random encounters. 


Study Area 

The study area is located in the foothills of west-cen- 
tral Alberta, near the town of Grande Cache (54°N 
119°W). The area is classed into subalpine and boreal 
natural subregions (Beckingham and Archibald 1996), 
and contains several main rivers and a dendritic pattern 
of creeks; lakes are scarce. Elevations range from 1300- 
1800 metres, and the climate is subarctic, with short 
wet summers and long cold winters. Temperatures aver- 


one 


574 


age 16°C in July and -13.5°C in December (Becking- 
ham and Archibald 1996). The forests are primarily 
Lodgepole Pine (Pinus contorta) and some White 
Spruce (Picea glauca). The wetland complexes support 
mostly Black Spruce (Picea mariana) and some Tama- 
rack (Larix laricina). Some south facing slopes support 
Trembling Aspen (Populus tremuloides) and willow 
(Salix sp.). 

This area supports a high diversity of large mam- 
mals: Woodland Caribou, Moose, Elk (Cervus elaphus), 
White-tailed Deer, Mule Deer (Odocoileus heminous), 
Bighorn Sheep (Ovis canadensis), Mountain Goats 
(Oreamnos americanus) and wild Horses (Equus caba- 
lus). Wolves, Coyotes (C. latrans), Grizzly Bears (Ursus 
arctos), Black Bears (Ursus americanus) and Cougars 
(Felis concolor) also exist throughout the study area. 

Major land use activities include forest harvesting, 
oil and gas exploration and development, coal mining, 
commercial trapping, and public uses such as hunting, 
fishing, hiking, horse packing and camping. Access is 
primarily on roads created for resource extraction, 
pipelines and seismic lines. Further descriptions of the 
study area can be found in Smith et al. (2000). 


Wolf captures and radio-tracking 

In January 2000, three Wolves from different packs 
(Simonette, Cutbank and Prairie Creek) were captured 
and immobilized by helicopter darting (Ballard et al. 
1991) or netgunning (Kuzyk 2002) and instrumented 
with GPS radiocollars (Lotek Engineering Sytems, 
Newmarket, Ontario). All Wolf handling was approved 
by the University of Alberta’s Faculty of Agriculture, 
Forestry and Home Economics Animal Care Policy 
(Number 96-99D), subject to the protocols of the 
Canadian Council of Animal Welfare. GPS collars were 
programmed to take one location per hour. 

During a short period of intensive sampling, from 2- 
15 March 2000, these radiocollared Wolves and their 
associated pack members were followed by radiotrack- 
ing from an airplane (Mech 1974). Wolves were relo- 
cated twice daily in hopes of detecting Wolf-killed 
deer (Fuller 1989). When a Wolf pack was located, the 
Wolves were counted and the area searched for ungu- 
late carcasses. If an ungulate kill or most Wolf pack 
members were not immediately found, Wolf trails were 
backtracked until an ungulate carcass was found (Hayes 
et al. 2000). 


Wolf kill sites 

An ungulate kill was assumed to be caused by 
Wolves if there was evidence of bloodstained snow, a 
disarticulated carcass and Wolf trails indicating a suc- 
cessful chase (Hayes et al. 2000). Wolves were assumed 
to be scavenging if the carcass was on its sternum 
(Ballard et al. 1987) or human sign indicated the ungu- 
late had been shot or road-killed. A GPS location was 
taken from an aircraft when directly over the kill site, 
with an estimated error of 73-128 metres (Carrel et al. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


1997). Kill sites were visited twice daily until the 
Wolves abandoned the carcass. Wolves were classified 
as being near a kill when all or most members of the 
pack were seen within one kilometre of the new kill site. 

Dead Moose were classified from the air as adult 
or calf (Peterson 1977). The amount of meat removed 
from the carcass was estimated (Carbyn 1983) and the 
number and behaviour of Wolves were recorded. All 
Wolf-killed ungulates for which species, sex and age 
(adult-calf) could not be confirmed from an airplane, 
and all others that were easily accessible with a heli- 
copter, were later ground-inspected. 


Data analysis 

GPS location data were differentially corrected using 
N4Win Version 2.40, which reduced location error to 
4-5 metres (Rempel and Rodgers 1997). Median travel 
distances in metres per one-hour interval (m/hr) were 
classified as kill site or non-kill site by calibrating the 
GPS data with field observation data. This was achieved 
by establishing a median time between each aerial 
observation in which Wolves were either near or away 
from an ungulate kill site. For example, if a Wolf pack 
was observed travelling at 1800 hr in the evening, and 
then relocated near a recent ungulate kill at 0800 hr the 
following morning (a time of 14 hours), the travel dis- 
tances in the first seven-hour period were placed in the 
non-kill site category. Those distances in the remaining 
seven hour period were placed in the kill site category. 
While both calculation of straight-line distances between 
remotely-collected locations and proportional assign- 
ment of time intervals to behaviour categories can intro- 
duce errors in absolute representation of patterns, asso- 
ciated errors were consistent across all data collected. 
Further, because we were interested in relative patterns, 
rather than absolute measures, we do not feel these lim- 
itations unduly compromise our results. 

All data were tested for normality before analysis 
and non-parametric procedures were used. All analyses 
were completed using SYSTAT (Version 8.0, SPSS Inc. 
1998). To determine if travel distances differed when 
Wolves were near or away from ungulate kill sites, the 
GPS locations from the collared Wolves were pooled 
and classified into two categories: at or away from an 
ungulate kill site. A Mann-Whitney U-test was used to 
test for differences in travel distance between these two 
categories. To examine variation in travel rates among 
wolf packs, location data for each pack were similarly 
divided into two categories (kill site/non-kill site), and 
analysed separately using Mann-Whitney U-tests. 


Results 

The three Wolf packs were located at seven ungulate 
kill sites from 2-15 March 2000 (Table 1). The Simon- 
ette Wolf pack made multiple kills at two sites: a cow 
and calf Moose were killed within 500 metres of each 
other on or near the same day, and the scattered remains 
of two deer kills were found within 100 metres of each 


2005 


KUZYK, ROHNER, and SCHMIEGELOW: TRAVEL RATES OF WOLVES 


575 


TABLE |. Ungulate kill sites of three Wolf packs during 2-15 March 2000 in west-central Alberta. 


Wolf Pack and Size (n) Adult Moose 
Cutbank (n = 8) | 
Prairie Creek (n = 5) 0 
Simonette (n = 11) ex 
Totals 4 


Calf Moose Adult Elk Deer 
| 0 0 
0 | 0 
0 0 
| | 1 


“includes one multiple kill of 1 cow and | calf Moose which is considered | kill 


* 


“includes one multiple kill of 2 deer which is considered | kill 


TABLE 2. Wolf travel distances (m/hr) at and away from seven ungulate kill sites as determined by one-hour GPS locations 
from three Wolves in separate packs during 2-15 March 2000 in west-central Alberta. 


All Travel 
(m/hr) 
Number of cases 553 
Minimum 0.2 
Maximum 6100 
Standard Deviation 749 
Median* 80 


“(Mann-Whitney U test P < 0.001) 


other (Table |). To be conservative, multiple kills were 
pooled for analysis. 

Wolves travelled a median distance of 80 m/hr dur- 
ing 14 consecutive days of monitoring (Table 2). They 
moved a median distance of 45 m/hr when near ungu- 
late kill sites, which differed significantly from a medi- 
an distance of 190 m/hr per hour when they were not 
near kill sites (U = 26 362, P< 0.001). 

Patterns in travel distance varied among Wolf packs. 
The Cutbank Wolf pack, which was only observed on 
Moose kills during the monitoring period, showed a 
highly significant difference (U = 1667, P < 0.001) in 
travel distances when at or away from kill sites. The 
Simonette pack, which was found near both Moose and 
deer kills, showed a marginally significant difference 
(U = 2740, P = 0.067) between travel distances at and 
away from kill sites. During aerial monitoring, only 
one Elk kill was recorded for the Prairie Creek pack 
and no difference (U = 2702, P = 0.601) in travel dis- 
tances related to kill sites was detected (Table 3). 


Discussion 

In this study, GPS radiocollar technology allowed 
Wolf travel rates to be recorded on a continuous (hourly) 
basis, irrespective of daylight and weather, and account- 


Travel at Kill Travel Away from Kill 


(m/hr) (m/hr) 
288 265 
0.2 0 
2044 6100 
326 963 
45 190 


ed for associated feeding, resting and other social behav- 
iours. Further, by combining GPS radio-collar technol- 
ogy with traditional methods used to study wolf kill 
rates (Mech 1974), a more comprehensive representa- 
tion of wolf travel rates was established. Results from 
this approach found a clear difference in Wolf travel 
rates related to ungulate kill sites, which is consistent 
with both anecdotal information and other research 
(Mech 1966; Peterson 1977; Hayes et al. 2000). 

We found that Wolves travelled a median distance of 
0.08 km/hr, which is substantially lower than that report- 
ed for Wolves travelling in the forest during winter 
(1.6-6.1 km/hr) (Musiani et al. 1998), on iced surfaces 
(8 km/hr) (Mech 1966), and on tundra during summer 
(8.7 km/hr) (Mech 1994). This difference could be large- 
ly due to the advantages of continuous (hourly) GPS 
monitoring in this study. While the data-collecting peri- 
od seldom lasted more than a few hours in other stud- 
ies, due to the requirement for maintenance of visual or 
auditory contact with the Wolves (Musani et al. 1998; 
Mech 1994), GPS technology provides for continuous 
data collection. As such, it provides information on 
movement patterns of Wolves that includes time spent 
in a variety of behaviours, such as resting, and stopping 
at old kill sites. 


TABLE 3. A comparison of median Wolf travel distances (m/hr) at and away from ungulate kill sites (n = number of GPS 
locations) for three Wolves in separate packs during 2-15 March 2000 in west-central Alberta. 


Wolf Pack Travel at Kill Travel Away from Kill Mann-Whitney 
(m/hr) (m/hr) U test (P) 
Cutbank 37 @= 135) 148 (n=48)s <0.001 
Prairie Creek 357 (n= 36) 338° (a= 159) 0.601 
Simonette 37 a = 016) a7 ae ='57) 0.067 


576 


However, these results are also lower than those 
reported by James (1999) for a boreal region of Alberta, 
where GPS collared Wolves moved 0.5 km/hr in the 
forest and averaged 1.4 km/hr on linear corridors. James 
(1999) collected GPS locations every five minutes with 
the objective of establishing Wolf speed, whereas in 
this study the collars obtained hourly locations with the 
purpose of establishing coarser estimates of Wolf travel 
distances in relation to kill sites. Frequency of locations 
can influence estimation of travel rates due to the linear 
extrapolation required between time intervals. In addi- 
tion, the boreal region of Alberta has less topographic 
relief than the foothills of west-central Alberta, which 
could also account for some difference in Wolf travel 
rates. Finally, prey type and density, as well as pack and 
territory size could also influence movement patterns 
and resultant estimates of travel rates. Further investi- 
gation of these differences is warranted in order to bet- 
ter understand regional patterns. 

The travel rate of Wolves in relation to ungulate kill 
sites provides important information when assessing 
predation risk to Caribou. Wolf travel distances were 4.2 
less when near kill sites (45 m/hr) than when away from 
kill sites (190 m/hr). We suggest that differences in trav- 
el patterns between the three Wolf packs studied were 
likely due to the different prey species each pack was 
hunting. The Cutbank pack showed the greatest differ- 
ences in travel at and away from kill sites and was found 
only at Moose kills. In one case, the pack remained near 
a cow Moose carcass for longer than four days, which 
reduced overall travel during the sampling interval. 

The Simonette Wolf pack travelled marginally shorter 
distances when at, compared with away from ungulate 
kills. Their travel near kills was, in fact, the same as the 
Cutbank pack, but they travelled much less when away 
from kill sites. This result may be due to a combination 
of an overall high kill rate of ungulates (Kuzyk 2002) 
and partial consumption of prey (Carbyn 1983). As 
well, this Wolf pack made multiple kills of ungulates 
(e.g., Ballard et al. 1987; Mech et al. 1998) which would 
reduce their overall travel. 

The Prairie Creek pack showed no difference in 
travel related to ungulate kills, with only one Elk kill 
being documented during aerial monitoring. This Wolf 
pack contained the fewest members (n = 5) during the 
study and pack size can affect ungulate kill rates (Fuller 
1989; Schmidt and Mech 1997; Hayes et al. 2000). 
Nevertheless, it also likely that the Prairie Creek pack 
was preying on deer, as these Wolves were observed 
hunting deer on a number of occasions, although no 
deer kills were found (see also Carbyn 1974). Fuller 
(1989) discusses in detail the logistical problems of 
determining Wolf kill rates of deer, due to the short 
time frame in which Wolves handle deer carcasses and 
the difficulty in detecting Wolf-killed deer from the 
air. [f deer are the main prey for the Prairie Creek pack, 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


then Caribou may face greater predation risk in this ter- 
ritory, than in the territories of packs preying primarily 
on Moose, due to associated increases in travel and 
encounter rates. The extensive, recent logging in the 
area (Smith et al. 2000) could favour deer numbers, 
and thereby influence predation risk to Caribou. Wolves 
in this study area were found to use forest cutblocks 
proportionately more than unharvested forests, which 
may be a response to increased number of deer and 
other ungulates that are attracted to the young vegeta- 
tion in the cutblocks (Kuzyk et al. 2004). A greater 
understanding of the role of deer in this Wolf-prey sys- 
tem is required, as most of the forest in westcentral 
Alberta has been allocated for timber harvest. 
Projecting the long-term implications of ongoing 
development activities for Wolf-prey systems requires 
a more detailed understanding of the responses of all 
species to landscape change (Bergerud 1974), and both 
the functional and numerical response of Wolves to 
changes in ungulate prey abundance and distribution at 
relevant spatial scales (see Lessard 2005). The results 
of this study provide information on movement pat- 
terns of Wolves in relation to ungulate kills, which 
could be expanded by calibrating the more extensive 
GPS-database with signals detected from concurrent 
aerial observations (Franke 2004). This would result in 
a larger sample of potential kill sites for further analy- 
sis of predation patterns in relation to landscape features. 


Acknowledgments 

Funding for this research was provided by the West- 
Central Alberta Caribou Standing Committee, the 
Alberta Sport, Recreation, Parks and Wildlife Founda- 
tion, and a University of Alberta Challenge Grant in 
Biodiversity (supported by the Alberta Conservation 
Association). A University of Alberta Graduate Re- 
search Assistantship and the Margaret Brown Award in 
Environmental Studies and Wildlife Resources provid- 
ed financial assistance to G. Kuzyk. We acknowledge 
the safe piloting conducted by Clay Wilson from Big- 
horn Helicopters and Denny Dennison from Coyote 
Air during wolf captures and monitoring. 


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Notes 


Caching Behavior by Wintering Northern Saw-Whet Owls, 


Aegolius acadicus 


ARNOLD DEVINE! and DwiGHuTt G. SMITH? 


'18 South Street, Plymouth, Connecticut 06782 USA 


"Biology Department, Southern Connecticut State University, New Haven, Connecticut 06515 USA 


Devine, Arnold, and Dwight G. Smith 2005. Caching behavior in Northern Saw-whet Owls, Aegolius acadicus. Canadian 


Field-Naturalist 119(4): 578-579. 


We observed 16 instances of caching behavior by Northern Saw-whet Owls in southern Connecticut between 30 October 
and 29 March over a 23-year period 1982 to 2004. Caches consisted of a single prey item in 13 instances and two prey items 
in three instances. Prey was cached either directly beneath the owl or from 5-28 cm distant, always on the same branch on 
which the owl was roosting. Observations of cached prey marked in the morning suggested that it was consumed after 14:30 
in the afternoon of the same day. Such a delay indicated a true cache rather than delayed feeding. 


Key Words: Northern Saw-whet Owl, Aegolius acadicus, caching behavior, Connecticut. 


A cache is defined as a hiding place or storage site, 
particularly of food. In birds, caching is the act of plac- 
ing excess food in storage for future consumption. 
Caching has been described for a number of species 
and is especially prevalent in certain raptors. We have 
observed caching behavior in Boreal Owls (Aegolius 
funereus), Eastern Screech-Owls (Megascops asio), 
Great Horned Owls (Bubo virginianus), and Northern 
Saw-whet Owls (personal obervations). In this paper 
we detail aspects of Northern Saw-whet Owl caching 
behavior that we observed at migratory and winter 
roosts in Connecticut in 1982-2004. 

Northern Saw-whet Owl caching was noted by 
Cannings (1993), Catling (1972), and Bent (1938). 
Bondrup-Nielsen (1977) described thawing of frozen 
prey by captive Saw-whet Owls and discussed this 
behavior in relation to caching. Bent (1938) related 
Saw-whet Owl caching behavior noted by Bendire 
(1877), who stated that he fed several whole bird car- 
casses to a captive Saw-whet Owl which immediately 
ate their heads, afterwards covering the bodies with 
loose feathers in the corner of its cage. 

We have previously described ecology and food 
habits of wintering Northern Saw-whet Owls in Con- 
necticut (Devine and Smith 1994; Smith and Devine 
1982); food consisted primarily of small mammals, 
mainly woodland mice (Peromyscus sp.), House Mouse 
(Mus musculus), jumping mice (Zapus sp.), shrews 
(Blarina brevicauda and Sorex sp.), chipmunks (Jamias 
striatus), and a variety of small birds, mostly species 
of sparrows and juncos. 

We observed 16 instances of this caching behavior 
(Table 1) from 30 October to 29 March between 1982 
and 2004. Each cache consisted of a single prey item 


in 13 instances and two prey items in three instances. 
Cached prey were all small animals, mostly rodents 
which is consistent with the Saw-whet Owl’s reputa- 
tion as a mouse predator. The majority (11 of 19 prey 
items or 57.9%) of cached prey were White-footed 
Mice (Peromyscus leucopus), but we also observed two 
House Mice, one Short-tailed Shrew, one Song Spar- 
row (Melospiza melodius), and one Pickerel Frog (Rana 
palustris), the last in October. 

Cached prey were laid across twigs and branchlets 
beneath the owl’s talons or within 5-28 cm. Most cached 
prey was within 5 cm of the roosting owl. The caching 
behavior that we observed is unlike that in other owls, 
which usually have a definite cache site. Instead, it 
sometimes resembles caching behavior exhibited by 
shrikes (Lanius species). To determine if these repre- 
sented food caches or were simply prey to be consumed 
shortly, one of us marked prey (n = 6) on branches 
with food coloring dyes when first seen, generally at 
07:00-09:00 hours and then checked the prey items 
in the afternoon of the same day, at 12:00-14:30 hours. 
In six instances, none of the food items had been con- 
sumed in these intervals. Follow-up visits the next day 
or two days later revealed that cache items were gone, 
presumably having been consumed either in the late 
afternoon or during that night. 

At least 14 of the 19 prey were decapitated when 
first observed. Neither the unidentified sparrow nor 
frog had been decapitated and we were unable to deter- 
mine the status of the three cached mammals because 
their anterior parts were partly covered by the owl’s 
feathers. 

Catling (1972) commented on caching by migrant 
Saw-whet Owls in Ontario. He suggested that owls 


578 


2005 


NOTES 


af9 


TABLE |. Cached prey of Northern Saw-whet Owls in Connecticut, 1982-2004. 


Species October 


White-footed Mouse (Peromyscus leucopus) 

House Mouse (Mus musculus) ] 
Short-tailed Shrew (Blarina brevicauda) 
Masked Shrew (Sorex cinereus) 
Golden-crowned Kinglet (Regulus satrapa) 
Song Sparrow (Melospiza melodia) 
Unidentified Sparrow 

Pickerel Frog (Rana palustris) 

Totals 2 


— 


consumed the heads of their prey during the previous 
night’s hunting episode, and the body sometime dur- 
ing the following day. Our observations substantiate 
this possibility but suggest that the remaining prey 1s 
consumed much later the following day (after 14:30) 
and thus represents a true cache. We also note a distinct 
relationship between weather and Saw-whet Owl 
caching; caching was most commonly observed (13 of 
16 instances) during the coldest months of the year, 
usually associated with periods of prolonged snow 
cover extending for 1-5 weeks. 

Catling (1972) reported 31 instances of caching; 
25 caches (80.6%) were during the migration period 
(defined as 1 October to 15 November and 15 March to 
30 April) and six (19.4%) from the winter (15 Novem- 
ber to 15 March. All 31 caches were decapitated and 
mice typically had forelimbs removed while birds had 
wings eaten. Our results differ markedly from Catling’s. 
Using his criteria for winter and migration periods, we 
found 14 caches (87.5 %) during the winter and two 
caches (12.5 %) from the migration period. Further- 
more we recorded two prey items at three different 
cache sites whereas this event went unreported by 
Catling. Thus caching appears to be an important sur- 
vival mechanism of wintering Northern Saw-whet Owl 
in southern Connecticut. 


January February March Totals 
5 5 | 11 
| 2 
l | 
| | 
| | 
| | 
| l 
l 
6 9 2 19 


Literature Cited 

Bendire, C. 1877. Notes on some of the birds found in 
southeastern Oregon, particularly in the vicinity of Camp 
Harney, from November, 1874, to January, 1877. Proceed- 
ings of the Boston Society of Natural History 19: 109.149. 

Bent, A. C. 1938. Life Histories of North American Birds of 
Prey. Part 2. Orders Faconiformes and Strigiformes, Smith- 
sonian Institution, United States National Museum Bul- 
letin (170). 482 pages. Washington, D. C. 

Bondrup-Nielsen, S. 1977. Thawing of frozen prey by bore- 
al and saw-whet owls. Canadian Field Naturalist 90: 477- 
479. 

Cannings, R. J. 1993. Northern Saw-whet Owl. The Birds of 
North America (42). Edited by A. Poole, P. Stettenheim, 
and F. B. Gill. Philadelphia Academy of Natural Sciences. 
Washington, D. C. 

Catling, P. M. 1972. Food and pellet analysis studies of the 
Saw-whet Owl (Ageolius acadicus. Ontario Field Biologist 
26: 72-85. 

Devine, A., and D. G. Smith. 1994. Northern Saw-whet Owl. 
Pages 174-175 in The Atlas of Breeding Birds of Connecti- 
cut. Edited by L. R. Bevier. State Geological and Natural 
History Survey of Connecticut Bulletin (113). 461 pages. 

Smith, D. G., and A. Devine. 1982. Winter food of the Saw- 
whet Owl. Connecticut Warbler 2: 52-54. 


Received 28 July 2004 
Accepted 15 September 2005 


580 THE CANADIAN FIELD-NATURALIST Vol. 119 


First Confirmation of Cougar, Puma concolor, in the Yukon 


THOMAS S. JUNG and PHILIP J. MERCHANT 


Yukon Department of Environment, Fish and Wildlife Branch, Box 2703, Whitehorse, Yukon, Y1A 2C6 Canada; e-mail: 
thomas.jung @ gov.yk.ca 


Jung, Thomas S., and Philip J. Merchant. 2005. First confirmation of Cougar, Puma concolor, in the Yukon. Canadian Field- 
Naturalist 119(4): 580-581. 


Cougar (Puma concolor) have been reported from the Yukon as early as 1944. Despite many sightings, no indisputable, 
physical evidence of Cougar being present in the Yukon had been obtained. Here, we report on the first. In November 2000, 
a specimen was secured from near Watson Lake, in southeastern Yukon. Whether this specimen, and the numerous sighting 


records, are indicative of a low-density breeding population in the Yukon, or represent transients, is unknown. 


Key Words: Cougar, Puma concolor, Puma, Mountain Lion, Yukon. 


The Cougar (Puma concolor) is one of the most 
widespread terrestrial mammals in the Americas, with 
their reported range extending from southeastern Alas- 
ka to Patagonia (Logan and Sweanor 2000). Globally, 
Cougars are classified by the IUCN (World Conserva- 
tion Union) as Near Threatened, thus they require 
careful attention by wildlife managers. In the Yukon, 
Cougars appear to have naturally occurred only in 
recent history (<100 years), similar to other species 
such as Coyote (Canis latrans), Mule Deer (Odo- 
coileus hemionus), and White-tailed Deer (O. virgini- 
anus; Youngman 1975; Hoefs 2001). The first reported 
Cougar sighting in the Yukon was in 1944. Since then, 
reported sightings, of variable reliability, have increased 
steadily. Despite local knowledge and expert opinion 
suggesting that Cougar were present in the Yukon, no 
physical evidence (e.g., scat, hair, confirmed tracks, 
photographs, or a specimen) had been found to sub- 
stantiate any sightings. Here, we report the first 
irrefutable evidence of Cougar in the Yukon. 

On 12 November 2000, a dead Cougar was discov- 
ered approximately 3 km NE of Watson Lake, Yukon 
(60.06°N, 128.70°W) by two local residents (E. 
Murphy-Kelley and J. Rhodes). The Cougar was 
found in a derelict car; apparently the abandoned car 
was being used as shelter by the Cougar. Interestingly, 
it appeared from tracks in the snow that there was 
another Cougar traveling with the one found. The 
Cougar appeared to have died shortly before being dis- 
covered, as it was not yet fully frozen despite tempera- 
tures being about -8°C. 

The specimen was retrieved, measured, aged, and 
necropsied. Standard morphometric measurements 
were made using established protocols for Cougars (I. 
Ross personal communication) while the carcass was 
laid laterally. Skull measurements were taken from a 
cleaned skull. Aging was done by gumline recession 
(Laundré et al. 2000). A field necropsy was performed 
to determine the cause of death, and select organs and 
tissues were examined at the Western College of Vet- 
erinary Medicine (Saskatoon, Saskatchewan) for dis- 


ease and other disorders. The specimen’s hide was pre- 
pared as a mount and it is now in the collections of 
the MacBride Museum (Whitehorse, Yukon); the skull 
and skeleton reside with the Yukon Department of 
Environment (Whitehorse, Yukon). 

The Cougar was an adult male that was estimated at 
>3 years old, and appeared normal except that it was 
emaciated — it weighed only 37.7 kg (males >2 years 
old normally weigh >53 kg; Logan and Sweanor 
2000). Morphometric measurements were: total length 
= 223 cm; tail lensth = 83 cm; chest girth = Gi cm; 
high neck circumference = 40 cm; maximum neck cir- 
cumference = 42.5 cm; right hind pad length = 41 mm; 
right hind pad width = 49 mm; head circumference = 
50.5 cm. Skull measurements were: length = 217 mm; 
zygomatic breadth = 177 mm. No disease or other dis- 
orders were apparent. The stomach was empty and it is 
believed that the cause of death was starvation. 

At the periphery of the Cougar’s range, sightings 
are normally all that is available to judge whether a 
low-density or transient population of Cougars exists 
(e.g., Gerson 1988; Cumberland and Dempsey 1994; 
Stocek 1995; Gau et al. 2001), and this evidence, on 
its own, is inconclusive. Finding this specimen con- 
firms that Cougars are occasionally found in the 
Yukon. Records also exist from adjacent jurisdictions: 
a specimen was obtained in Wrangell, Alaska, in 1989, 
there are an increasing number of sighting records in 
eastern and southeastern Alaska (Alaska Geographic 
Society 1996), and animals have been sighted (but 
with no corroborating evidence) west of the Macken- 
zie River in the Northwest Territories (Gau et al. 
2001). Six unconfirmed Yukon sighting records in- 
clude kittens (Yukon Department of Environment, 
unpublished data), suggesting a breeding population. 
Alternatively, Cougars seen in the Yukon may repre- 
sent a few individuals (e.g., young males) that make 
long distance dispersals and are not indicative of a 
viable local breeding population (Pierce et al. 1999; 
Sweanor et al. 2000). In Wyoming, one radio-collared 
male Cougar is known to have dispersed >1000 km 


2005 


(Thompson and Jenks 2005). Regardless, Cougars 
appear to be a rare but regular component of the mam- 
malian fauna of the Yukon. 

This confirmation of the addition of Cougar to the 
list of species in the Yukon is representative of the rel- 
atively dynamic state of the mammalian fauna of 
northern biomes. Other northern areas are also wit- 
nessing colonization by some species of large mam- 
mals. For example, Moose (Alces alces) and Coyotes 
are relatively new additions to the mammalian fauna 
of Labrador (Chubbs and Schaefer 1997; Chubbs and 
Phillips 2002). In the Northwest Territories, White- 
tailed Deer and Cougar also appear to be expanding 
northward (Veitch 2001; Gau et al. 2001). 


Acknowledgments 

We are grateful to E. Murphy-Kelly and J. Rhodes 
for promptly reporting this Cougar to wildlife officials. 
R. Hennings and D. Rudd retrieved the carcass. A. 
Baer, M. Hoefs, D. Rudd, E. Neufeld, and T. Grabow- 
ski assisted in processing and preserving the speci- 
men. M. Oakley, D.V.M., undertook the necropsy and 
F. Leighton, D.V.M., tested the organs for disease and 
other disorders. D. Nagorsen, B. Slough, J. Adam- 
czewski, and R. Maraj kindly provided helpful com- 
ments on an earlier draft. We thank the late I. Ross for 
assisting us with measurement and aging protocols. 


Literature Cited 

Alaska Geographic Society. 1996. Mammals of Alaska. 
Alaska Geographic Society, Anchorage, Alaska. 176 
pages. 

Chubbs, T. E., and J. A. Schaefer. 1997. Population growth of 
Moose, Alces alces, in Labrador. Canadian Field-Natural- 
ist 111: 238-242. 

Chubbs, T. E., and F. R. Phillips. 2002. First record of an 
Eastern Coyote, Canis latrans, in Labrador. Canadian 
Field-Naturalist 116: 127-128. 


NOTES 581 


Cumberland, R. E., and J. A. Dempsey. 1994. Recent con- 
firmation of a Cougar, Felis concolor, in New Bruns- 
wick. Canadian Field-Naturalist 108: 224-226. 

Gau, R. J., R. Mulders, T. Lamb, and L. Gunn. 2001. 
Cougars (Puma concolor) in the Northwest Territories 
and Wood Buffalo National Park. Arctic 54: 185-187. 

Gerson, H. B. 1988. Cougar, Felis concolor, sightings in 
Ontario. Canadian Field-Naturalist 102: 419-424. 

Hoefs, M. 2001. Mule, Odocoileus hemionus, and White- 
tailed, O. virginianus, Deer in the Yukon. Canadian 
Field-Naturalist 115: 296-300. 

Laundré, J. W., L. Hernandez, D. Streubel, K. Altendorf, 
and C. L. Gonzalez. 2000. Aging mountain lions using 
gum-line recession. Wildlife Society Bulletin 28: 963- 
966. 

Logan, K. A., and L. L. Sweanor. 2000. Puma. Pages 347- 
377. In Ecology and management of large mammals in 
North America. Editied by S. Demarais and P. Kraus- 
man. Prentice-Hall, Englewood Cliffs, New Jersey. 

Pierce, B. M., V. C. Bleich, J. D. Wehausen, and R. T. 
Bowyer. 1999. Migratory patterns of Mountain Lions: 
implications for social regulation and conservation. Jour- 
nal of Mammalogy 80: 986-992. 

Stocek, R. F. 1995. The Cougar, Felis concolor, in the Mar- 
itime provinces. Canadian Field-Naturalist 109: 19-22. 
Sweanor L. L., K. A. Logan, and M. G. Hornocker. 2000. 
Cougar dispersal patterns, metapopulation dynamics, and 

conservation. Conservation Biology 14: 798-808. 

Thompson, D. J., and J. A. Jenks. 2005. Long-distance dis- 
persal by a subadult male Cougar from the Black Hills, 
South Dakota. Journal of Wildlife Management 69: 818- 
820. 

Veitch, A. M. 2001. An unusual record of a White-tailed 
Deer, Odocoileus virginianus, in the Northwest Territories. 
Canadian Field-Naturalist 115: 172-175. 

Youngman, P. M. 1975. Mammals of the Yukon Territory. 
National Museums of Canada, Ottawa, Ontario. 192 


pages. 


Received 2 February 2004 
Accepted 19 September 2005 


582 THE CANADIAN FIELD-NATURALIST Vol. 119 


First Occurrence of the Round Goby, Neogobius melanostomus, 
in the St. Lawrence River at Cornwall, Ontario 


M. BRIAN C. HICKEY! and ADRIENNE R. FOWLIE? 


'St. Lawrence River Institute of Environmental Sciences, Windmill Point, 2 Belmont Street, Cornwall, Ontario K6H 4Z1 
Canada; e-mail: bhickey @riverinstitute.ca 
*Queen’s University, Kingston, Ontario K7L 3N6 Canada; e-mail: fowliea@biology.queensu.ca 


Hickey, M. Brian C., and Adrienne R. Fowlie. 2005. First occurrence of the Round Goby, Neogobius melanostomus, in the 
St. Lawrence River at Cornwall, Ontario. Canadian Field-Naturalist 119(4): 582-583. 


We document the first reported occurrence of the Round Goby, Neogobius melanostomus, a small benthic fish native to the 
Black and Caspian seas, in the St. Lawrence River near Cornwall. On 7 September 2004, we observed approximately 20 
Round Gobies while SCUBA diving at a depth of 7 m, downstream of the Saunders Generating Station at Cornwall, Ontario. 
Round Gobies appear to have arrived recently in this reach of the river and have not previously been detected despite exten- 


sive fish surveys conducted in the area. 


Key Words: Round Goby, Neogobius melanostomus, exotic species, St. Lawrence River, Cornwall, Ontario. 


The Round Goby, Neogobius melanostomus, is a 
small benthic fish native to the Black and Caspian 
seas (Charlebois et al. 2001). It was introduced to the 
Great Lakes from Eurasia in ballast water of trans- 
oceanic ships (Charlebois et al. 2001). The first report- 
ed sighting of Round Goby was in Lake St. Clair in 
1990, and within five years, the species was reported in 
all five Great Lakes (Charlebois et al. 2001). Reasons 
for this rapid expansion include its high fecundity, wide 
tolerance of abiotic factors, and broad diet (French and 
Jude 2001; Corkum et al. 2004). 

The Round Goby’s expansion in to the Great Lakes 
has been rapid but movement into the St. Lawrence 
River has been much slower; perhaps an indication 
that habitat or hydraulic conditions are sub-optimal in 
the St. Lawrence. Gobies have been recorded upstream 
of the Moses-Saunders Dam as close as Prescott, 
Ontario (approximately 100 km upstream) and in the 
lower St. Lawrence near Quebec City. The geographic 
distribution of the Round Goby suggests that there have 
been at least two separate introductions but we are not 
aware of any studies that confirm this view. To our 
knowledge, they have not been previously recorded in 
the Cornwall area. 

Their absence from the Cornwall area is probably a 
real phenomenon rather than the result of lack of 
detection. The St. Lawrence River between the Moses- 
Saunders Dam and the Beauharnois Dam in Quebec 
has been designated an Area of Concern (AOC) by 
the International Joint Commission. Consequently, 
the Cornwall area has been the focus of several ongo- 
ing biological investigations which have included 
regular surveys of the littoral zone fish community 
(Dreier et al. 1997; M. B. C. Hickey unpublished data). 
These surveys have included electrofishing, minnow 
trapping, and gill netting as well as snorkel and dive 
transects along an 8-km stretch of the St. Lawrence 
between the Moses-Saunders Dam and Windmill 
Point. 


On 7 September 2004, we observed approximately 
20 Round Gobies while SCUBA diving at a depth of 
7 m, downstream of the Saunders Generating Station 
at Cornwall, Ontario (45°00'31"’N, 74°46'09"). The 
Round Goby’s distinctive body shape and conspicu- 
ous dorsal fin spot (Figure 1) make it easy to distin- 
guish from other small benthic fish reported from the 
St. Lawrence River (e.g., darters, Etheostoma sp. and 
sculpins, Cottus sp.). On two dives, less than 3 km 
downstream of the site where we observed round gob- 
ies on 7 September 2004, we did not notice gobies, 
suggesting that the gobies we observed represent the 
leading edge of their expansion in the St. Lawrence 
River as they move downstream from the upper St. 
Lawrence River. 

The invasion of the Round Goby poses potential 
threats to native species such as the sculpins and darters 
that occupy similar habitat but are unable to compete 
for resources with the more aggressive gobies (Janssen 
and Jude 2001). The Round Goby’s use of zebra mus- 
sels as a major prey may dramatically alter the patterns 
of contaminant uptake in the St. Lawrence River food 
web. The potential role of sediment contamination in 
the Cornwall “Area of Concern” (Dreier et al. 1997) 
is the subject of several ongoing investigations (M. B. 
C. Hickey and A. Fowlie, unpublished data). The intro- 
duction of Round Goby and subsequent lengthening 
of the food chain has been associated with increased 
levels of PCBs in fish of higher trophic levels in west- 
ern Lake Erie (Morrison et al. 2000). The recent expan- 
sion of Round Goby in the St. Lawrence River AOC 
at Cornwall could produce similar effects in mercury 
biomagnification which has been shown to increase 
with food chain length (Cabana et al. 1994). Given the 
potential ecological and economic impacts of Round 
Goby invasion, the Cornwall area may provide a nat- 
ural laboratory in which to study the impacts of the 
Round Goby and ultimately devise mitigation or con- 
trol measures. 


2005 


NOTES 


FIGURE 1. Round Gobies, Neogobius melanostomus, captured in the St. Lawrence River, immediately downstream of the 
Saunders Generation Station at Cornwall, Ontario. Photo by M. B. C. Hickey and A. R. Fowlie. 


Acknowledgments 

We thank Jason Szwec, Rob Marion and Jessica 
Roy for assisting us in the field and Jason Szwec and 
Rob Gratton for reviewing an earlier version of the 
manuscript. Sean Hickey and Lisa Hickey assisted with 
the capture of the Round Gobies photographed for 
Figure 1. 


Literature Cited 

Cabana, G., A. Tremblay, J. Kalff, and J. B. Rasmussen. 
1994. Pelagic food chain structure in Ontario lakes: A 
determinant of mercury levels in Lake Trout (Salvelinus 
namaycush). Canadian Journal of Fisheries and Aquatic 
Sciences 51: 381-389. 

Charlebois, P. M., L. D. Corkum, D. J. Jude, and C. 
Knight. 2001. The round goby Neogobius melanostomus 
invasion: current research and future needs. Journal of 
Great Lakes Research 27: 263-266. 

Corkum, D. L., M. R. Sapota, and K. E. Skora. 2004. The 
round goby, Neogobius melanostomus, a fish invader on 


both sides of the Atlantic Ocean. Biological Invasions 6: 
173-181. 


Dreier, S. I., J. Anderson, J. Biberhofer, M. Eckersley, R. 
Helliar, M. B. C. Hickey, L. Richman, F. Stride, and 
The St. Lawrence Remedial Action Plan Public Advi- 
sory Committee. 1997. Great Lakes Great River: Reme- 
dial Action Plans for the St. Lawrence River (Cornwall) 
Area of.Concern. 203 pages. 

French, J. R. P., III, and D. J. Jude. 2001. Diets and diet 
overlap of nonindigenous gobies and small benthic 
native fishes co-inhabiting the St. Clair River, Michigan. 
Journal of Great Lakes Research 27: 300-311. 

Janssen, J. and D. J. Jude. 2001. Recruitment failure of 
mottled sculpin (Cottus bairdi) in Calumet Harbor, 
southern Lake Michigan, induced by the newly intro- 
duced round goby Neogobius melanostomus. Journal of 
Great Lakes Research 27: 319-328. 

Morrison, H. A., D. M. Hittle, and G. D. Haffner. 2000. 
The relative importance of species invasions and sedi- 
ment disturbance in regulating chemical dynamics in 
western Lake Erie. Ecological Modeling 125: 279-294. 


Received 7 February 2005 
Accepted 12 December 2005 


584 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Swift Fox, Vulpes velox, Den Located Next to a Railroad Track in 


Northwestern Texas 


Kerry L. NICHOLSON!: 2, BRADY K. MCGEE! 3, and WARREN B. BALLARD! 


'Department of Range, Wildlife, and Fisheries Management, Texas Tech University, Box 42125, Lubbock, Texas, USA 
Present address: School of Renewable Natural Resources, University of Arizona, 325 Biological Sciences East, Tucson, 


Arizona, USA 


Present address: Lower Rio Grande Valley National Wildlife Refuge, Rt. 2 Box 202-A, Alamo, Texas 78516 USA 


Nicholson, Kerry L., Brady K. McGee, and Warren B. Ballard. 2005. Swift Fox, Vulpes velox, dens located next to a rail- 
road track in northwestern Texas. Canadian Field-Naturalist 119(4): 584-585. 


Swift Fox (Vulpes velox) dens are typically found in areas were the vegetation is sparse, in loam soils, and with unobstructed 
views of the surrounding area. In 2002 a Swift Fox in northwest Texas was found in a unique den situated at the base of a 
hill with the entrance within Im of an active railroad track. Use of a den in such proximity to railroad tracks has never been 


previously reported. 


Key Words: Swift Fox, Vulpes velox, den, Texas. 


The Swift Fox (Vulpes velox) is one of the most 
burrow dependent canids in North America (Kilgore 
1969; Hines and Case 1991; Jackson and Choate 2000; 
Harrison 2003; Uresk et al. 2003). The objective of 
this paper is to describe an unusual den site used by a 
Swift Fox in northwest Texas. 

On 28 September 2002, we captured, radiocollared, 
and ear-tagged a juvenile female Swift Fox as part of 
a larger study on Swift Fox ecology (Nicholson 2004; 
McGee 2005). We monitored this fox until her death 
16 April 2003. This fox was occasionally located in 
dens with another resident adult male on our study site. 
She was killed by an automobile while moving between 
her den and our study site to the south. 

The female fox maintained five different den loca- 
tions including an unusual den located next to railroad 
tracks. We tracked this female to the den on three 
separate occasions between 25 March and 16 April 
2003. No other fox used this den. The railroad tracks 
cut through a hill with a den at the bottom of a 1-meter 
bank (Figure |). The den was underneath a limestone 
rock shelf 23 meters from Highway 54, and | meter 
from the railroad. Approximately 0.4 km west of the 
den was an active granary and train stop; to the east 
were rangeland and crop fields. The embankments on 
either side of the railroad tracks were covered in yucca 
(Yucca sp.). There were 20—25 trains per day that trav- 
eled along the tracks between Stratford and Dalhart, 
Texas (D. Richard, Union Pacific Railroad, personal 
communication). 

Visibility appears to be an important factor in den 
site and habitat selection for Swift Fox (Zoellick et al. 
1989; Uresk et al. 2003). Unlike most Swift Fox dens, 
which allow for a clear view of the surrounding area, 
this den had limited (i.e., | m) visibility. The other four 
dens this female used were typical of Swift Fox dens 
in short vegetation, loam and clay-loam soils, and were 
surrounded by heavily grazed rangeland. Vegetation 


surrounding the railroad den (yucca taller than a fox), 
poor visibility (due to steep embankments), proximity 
to the railroad, and placement in a limestone rock shelf 
(dry, rocky soil) was uncharacteristic of previously 
described Swift Fox den sites. 

We were not sure why this den site was selected but 
several explanations seem plausible. First, a lack of 
normal den sites or an over-saturation of foxes within 
the surrounding area may have limited this female’s 
choices. We documented 35 different dens sites used 
by 20 monitored foxes on our adjacent study site. Also, 
this unusual den could have been refuge from preda- 
tion. In our study site, Coyotes (Canis latrans) were 
heavily exploited by landowners and recreational hunt- 
ers, yet Coyotes were responsible for 80% of Swift Fox 
deaths (Kamler et al. 2003). If there was risk from 
Coyotes, having a den site next to the train tracks may 
have deterred pursuit by Coyotes. This particular den 
was located outside previously documented Coyote 
home ranges (Kamler et al. 2003). For a Swift Fox, 
danger from Coyotes may outweigh the danger from 
trains. The use of this den could have been at first op- 
portunistic shelter between divided range lands. It is 
probable that this den was constructed by some other 
animal like a Badger (Taxidea taxus) or Striped Skunk 
(Mephitis mephitis) and the fox came upon it while 
searching for other resources. This den was located 
slightly north of her natal home range and could have 
been a refuge den used in pre-dispersal exploration. 

Previous research suggested that Swift Fox dens 
tended to be near roads (Hillman and Sharps 1978; 
Hines and Case 1991; Pruss 1999). Kamler et al. (2003) 
found that Swift Foxes seemed naive to the threat of 
vehicles because they hunt in road ditches within a 
few meters of passing vehicles. Swift Fox mortalities 
due to vehicles suggested that foxes never learned to 
avoid vehicles (Kamler et al. 2004). Swift Foxes may 
regard trains in the same respect as vehicles, thus never 


2005 


NOTES 


585 


Figure 1. Picture of a swift fox den with an arrow indicating the entrance next to the railroad 
tracks in Sherman County, Texas, 2003. 


actively seeking to avoid trains. Avoidance of Coy- 
otes would be a reasonable hypothesis if Coyotes 
avoided railroad tracks. 


Acknowledgments 

This project was funded by the National Fish and 
Wildlife Foundation and Texas Tech University. We 
thank F. Pronger for allowing us to conduct studies 
on his land. We thank E. and B. Hampton for their 
endless support, and A. McGee and M. Green for 
data collection. Research protocols were approved by 
the Animal Care and Use Committee at Texas Tech 
University. This is Texas Tech University, College of 
Agricultural Sciences and Natural Resources Techni- 
cal Publication T-9-1000. 


Literature Cited 

Harrison, R. 2003. Swift fox demography, movements, 
denning, and diet in New Mexico. Southwestern Natural- 
ist 48: 261-273. 

Hillmann, C. N., and J. C. Sharps. 1978. Return of swift 
fox to northern Great Plains. Proceedings of South Dako- 
ta Academy of Science 57: 154-162. 

Hines, T. D., and R. M. Case. 1991. Diet, home range, 
movements, and activity periods of swift fox in Nebras- 
ka. Prairie Naturalist 23: 131-138. 

Jackson, V., and J. R. Choate. 2000. Dens and den sites of 
the swift fox, Vulpes velox. The Southwestern Naturalist 
45: 212-220. 


Kamler, J. F., W. B. Ballard, E. M. Gese, R. L. Harrison, 
S. Karki, and K. Mote. 2004. Adult male emigration 
and a female-based social organization in swift foxes, 
Vulpes velox. Animal Behavior 67: 699-702. 

Kamler, J. F., W. B. Ballard, R. L. Gilliland, and K. 
Mote. 2003. Spatial relationships between swift foxes 
and coyotes in northwestern Texas. Canadian Journal of 
Zoology 81: 168-172. 

Kilgore, D. L., Jr. 1969. An ecological study of the swift 
fox (Vulpes velox) in the Oklahoma panhandle. American 
Midland Naturalist 81: 512-534. 

McGee, B. K. 2005. Swift fox ecology in northwest Texas. 
Dissertation, Texas Tech University, Lubbock, Texas, USA. 

Nicholson, K. L. 2004. Swift fox occurrence in black tailed 
prairie dog towns in northwest Texas; with notes on 
movement. Thesis. Texas Tech University, Lubbock, 
Texas, USA. 

Pruss, S. D. 1999. Selection of natal dens by the swift fox 
(Vulpes velox) on the Canadian prairies. Canadian Jour- 
nal of Zoology 77: 646-652. 

Uresk, D. W., K. E. Severson, and J. Javersak. 2003. Veg- 
etative characteristics of swift fox denning and foraging 
sites in southwestern South Dakota. United States 
Department of Agriculture Forest Service, Rocky Moun- 
tatin Research Station, South Dakota, USA. 

Zoellick, B. W., N. S. Smith, and R. S. Henry. 1989. Habi- 
tat use and movements of desert kit foxes in western Ari- 
zona. Journal of Wildlife Management 53: 955-961. 


Received 29 March 2005 
Accepted 27 September 2005 


586 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Water-bears from the Rocky Mountains: A First Look at Alberta’s 


Tardigrade Fauna 


MATTHEW J. BOECKNER and HEATHER C. PROCTOR 


Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada; email: Boeckner@ualberta.ca 


Boeckner, Matthew J., and Heather C. Proctor. 2005. Water-bears from the Rocky Mountains: A first look at Alberta’s: tardigrade 


fauna. Canadian Field-Naturalist 119(4): 586-588. 


There is no information in the published literature regarding Alberta’s terrestrial water-bear (Tardigrada) fauna. We surveyed 
across an elevational gradient (1100 — 1800 ft) on Black Cat Mountain in the Alberta Rocky Mountain range. Ten taxa 
were identified from 902 specimens, all of which are new records for Alberta and one species is new for Canada: 
Macrobiotus cf. islandicus Richters, 1904. There was no clear relationship between elevation and tardigrade distri- 
bution, possibly owing to patchiness of these animals in the field and lack of adequate replication at each elevation. 


Key Words: Tardigrada, elevation diversity, faunistic survey, Black Cat Mountain, Alberta. 


Given the charismatic nature of water-bears (7ar- 
digrada) and relative ease of collection, it is remark- 
able that so little is known about the ecology of Cana- 
dian tardigrades (see Boeckner et al. 2005 for a review 
of studies on Canadian tardigrades). Species lists exist 
for some Canadian provinces and territories; however, 
most records were compiled during patchy trans-Cana- 
dian surveys conducted in the early 20" century 
(Richters 1908; Murray 1910). British Columbia, New- 
foundland and Labrador, New Brunswick, the Yukon 
and Northwest Territories have had the largest number 
of studies, British Columbia leading with five (Boeck- 
ner et al., 2005). Alberta, Manitoba, Nova Scotia and 
Prince Edward Island currently have no records of ter- 
restrial tardigrades; however, a single study identified 
members of the genus Hypsibius in five Albertan lakes 
(Anderson and DeHenau 1980). Also, Murray (1910) 
collected tardigrades from the Rocky Mountains; how- 
ever, it is unclear whether these samples were from 
Alberta or British Columbia. 

The aim of this study was to begin the first of 
many forays that will result in a comprehensive list of 
Albertan tardigrade fauna. Additionally, elevation has 
been investigated as a factor affecting tardigrade assem- 
blages (example: Guidetti et al. 1999). Thus, our data 
also provide information regarding distribution patterns 
across an elevational gradient. 


Materials and Methods 

The study was conducted on Black Cat Mountain 
(Figure 1) in the Rocky Mountain Foothills (Boule 
Range) east of Jasper National Park (53°18'N: 
117°52'W). Moss samples (~250 ml each) were col- 
lected from eight sites along an elevational gradient. 
The lowest altitude sampled was 1100 m, the highest 
was 1800 m and samples in between were collected 
at altitudinal intervals of 100 m. The moss collected 
was Pleurozium schreberi (Brid.) Mitt., which is a 
widespread species common in dry forested areas 
(Johnson et al. 1995). Two sub-samples were collect- 
ed at each altitude. Each of the 16 samples was placed 
in a separate paper bag for transport from the field. 


Tardigrade extraction from the moss was achieved 
by using a Baermann funnel apparatus (Whitehead and 
Hemming 1965). This involved placing a sample of 
moss wrapped in cheesecloth in the cone of a blocked 
funnel. The funnel was then filled with water to sub- 
merge the sample. Live tardigrades migrate through the 
cheesecloth and collect in the water at the narrow base 
of the funnel. After 48 hours, the water from the base 
of the funnel was collected by removing the stopper. 
Any tardigrades remaining in the moss were separated 
by agitating the sample repeatedly in water. The water 
from both of these techniques was then combined and 
examined under a stereomicroscope (minimum 25x) 
for tardigrades and their eggs. Specimens were slide- 
mounted using polyvinyl alcohol medium (commer- 
cially available as PVA from BioQuip Products Inc., 
catalogue number 6371). 

Identification of species was facilitated using the 
work of Ramazzotti and Maucci (1983). Hieronim 
Dastych (Zoologisches Institut und Zoologisches 
Museum, Universitaet Hamburg) and Nigel Marley 
(Faculty of Science, University of Plymouth) provided 
invaluable assistance by confirming the identities of 
voucher specimens. 


Results 

A total of 902 tardigrades representing seven gen- 
era and nine identifiable species were extracted from 
the 16 moss samples. Eighty-seven specimens were in 
poor condition and unidentifiable. Six of the species 
identified in this study (denoted by cf.) were from com- 
plexes composed of very similar sub-species (H. Dastych, 
personal communication). Tardigrade eggs are often 
required to identify sub-species within a complex. 
The tardigrades found in the greatest abundance were 
Ramazzottius sp. (N=247), Macrobiotus cf. hufelandi 
Biserov, 1991 (N=154), Macrobiotus cf. richtersi 
Murray, 1911 (N=107) and Milnesium tardigradum 
Doyere, 1840 (N=100). The Ramazzottius sp. may rep- 
resent more than one species; however, without eggs 
this could not be determined. The remaining species 
were each represented by fewer than 100 specimens: 


2005 


Hypsibius cf. convergens Urbanowicz, 1925 (N=79), 
Macrobiotus cf. areolatus Murray, 1907 (N=66), 
Macrobiotus cf. islandicus Richters, 1904 (N=29), 
Echiniscus spitsbergensis Scourfield, 1987 (N=23), 
Isohypsibius cf. tuberculatus Plate, 1888 (N=8) and 
Platycrista cheleusis Kathman, 1990 (N=2). 

All 10 taxa are new records for Alberta. One species 
was a new record for Canada (McInnes 1994; Boeckner 
et al., 2005): M. cf. islandicus. Platicrista cheleusis has 
only ever been recorded previously in British Colum- 
bia (Kathman 1990). Table 1 shows the distribution 
patterns of the nine tardigrades identified to species and 
locations in Canada where they have been previously 
recorded. 

An analysis of the distribution of species across an 
altitudinal gradient revealed only a few minor patterns. 
The third (1300 m) and fifth (1500 m) elevations con- 
tained the majority of specimens, 41% and 32%, 
respectively. Within these two altitudes Ramazzottius 
sp. dominated but was not found again within the other 
elevations. Milnesium tardigradum was relatively 
abundant in the third elevation but was not recorded 
elsewhere. Similarly, M. cf. islandicus and the one het- 
erotardigrade identified, E. spitsbergensis, were found 
only in the lower elevations (1100 to 1300 m). M. cf. 
hufelandi exhibited the most even distribution across 
the gradient. 


Discussion 

All 10 of the tardigrade taxa identified within this 
study were new records for Alberta and one species 
was a new record for Canada. The discovery of M. cf. 
islandicus was surprising given the limited distribution 
of this species reported in previous studies (Table 1). 
Ramazzotti and Maucci (1983) describe this species as 
neither frequently found nor hard to find suggesting 
that it may have a widespread distribution but generally 
low abundance. Platycrista cheleusis has previously 
found only on Vancouver Island, British Columbia 
(Kathman 1990). The remaining seven species are 


BOECKNER AND PROCTOR: ALBERTA’ S TARDIGRADE FAUNA 


587 


“ie 
Hey 


AINADA 


FiGurE |. Location of Black Cat Mountain (X) in the Rocky 
Mountain foothills just outside of Jasper National 
Park. 


reported to have widespread to cosmopolitan distribu- 
tions and therefore it is not surprising to identify them 
from Alberta mosses. It is important to note that 
although species designations are given here for most 
taxa, the lack of tardigrade eggs within the samples 
made some identification possible only to complexes 
of sub-species (denoted by cf.). Voucher specimens of 
each species were submitted to the University of Alber- 
ta freshwater invertebrate collection and are available 
upon request. 


Table 1. Nine species of tardigrades new to Alberta and where previously recorded in Canada (adapted with permission from 
Bateman and Collins 2001). Distribution patterns as noted by Ramazzotti and Maucci (1983) and McInnes (1994). Locations: 
BC, British Columbia; L, Labrador; NB, New Brunswick; NF, Insular Newfoundland: NWT, Northwest Territories; ON, 
Ontario; PQ, Province du Quebec. Recorded by: A, Argue (1971, 1972, 1974); B&C, Bateman and Collins (2001); B, Boeckner 
et. al. (2005); I, Iharos (1973); K, Kathman (1990); M, Murray (1910); P, Pilato (1977); R, Richters (1908); VR', Van Rompu 


et al. (1991); VR’, Van Rompu et al. (1992). 


Canadian Records 
NB (A), NWT (VR!7) 


Species 


Echiniscus spitsbergensis 
Macrobiotus cf. areolatus 
M. ct. hufelandi 

M. cf. islandicus 

M. cf. richtersi 

Aypsibius cf. convergens 
Isohypsibius cf. tuberculatus 
Platycrista cheleusis 
Milnesium tardigradum 


new Canadian record 
BC(K), NB(A), PQ(D), 


BC(M), NB(A) 
BC(K) 


BC(M,K), NB(A), ON(M), RM(M), SK(P) 
BC(R,M,K), L (B), NB(A), NF(B&C), ON(M), PQ(I), RM(M) 


BC(K), L (B), NB(A), NF(B&C), NWT(VR!”) 


BC (R, K), L (B), NB (A), NF(B&C), ON(M), PQ(D) 


Distribution 


widespread 
cosmopolitan 
cosmopolitan 
Europe-Iceland 
cosmopolitan 
cosmopolitan 
widespread 
Canada-BC 
cosmopolitan 


588 


The analysis of tardigrade distributions across an 
elevational gradient produced only weak patterns. This 
was likely owing to lack of adequate replication at each 
elevation and natural variation in tardigrade abundance 
and distribution (patchiness). Furthermore, sample sites 
were likely variable in more ways than elevation alone 
_ (.e., exposure, substrate type, moisture, etc.). Gener- 
ally, M. cf. hufelandi tended to have a more uniform 
distribution throughout most elevations when com- 
pared with the other species. Additionally, Echiniscus 
spitsbergensis and M. ct. islandicus had distributions 
limited to lower elevations. It remains to be determined 
whether these general trends are indicative of actual 
altitude-constrained distributions. 

Although this study has identified a fair number of 
Albertan tardigrades there are vast areas of the province 
that have yet to be surveyed. Until such investigations 
are made it is impossible to comment confidently on 
species ranges. 


Acknowledgments 

We thank Hieronim Dastych and Nigel Marley for 
lending their outstanding expertise in tardigrade identi- 
fication to this study. Much appreciation is also extend- 
ed to Kevin McEwan and Danica Belter who aided in 
collection and extraction of the samples. We gratefully 
acknowledge the financial support of this research by 
the Natural Sciences and Engineering Research Council 
Discovery Grant held by Heather C. Proctor. 


Literature Cited 

Anderson, R., and A. DeHenau. 1980. An assessment of the 
meiobenthos from nine mountain lakes in western Canada. 
Hydrobiologia 70: 257-264. 

Argue, C. W. 1971. Some terrestrial tardigrades from New 
Brunswick, Canada. Canadian Journal of Zoology 49: 401- 
415. 

Argue, C. W. 1972. Tardigrades from New Brunswick, Cana- 
da 2. Canadian Journal of Zoology 50: 87-94. 

Argue, C. W. 1974. Tardigrades from New Brunswick, Cana- 
da 3. Canadian Journal of Zoology 52: 919-922. 

Bateman, L. E., and M. Collins. 2001. A preliminary account 
of the tardigrades of Newfoundland. Zoologischer Anzeiger 
240: 223-232. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Boeckner, M., M. Collins, J. Finney-Crawley, and L. Bate- 
man. /n press. The bryofauna of remote coastal Labrador: 
including a review of current Canadian records. Zootaxa*. 

Guidetti, R., R. Bertolani, and D. R. Nelson. 1999. Ecologi- 
cal and faunistic studies on tardigrades in leaf litter of 
beech forests. Zoologischer Anzeiger, 238: 215-223. 

Iharos, G. 1973. Angaben zur geographischen Verbreitung 
der Tardigraden. Opuscula Zoologica (Budapest) 12: 73- 
86. 

Pilato, G. 1977. Macrobiotus willardi, a new species of tar- 
digrada from Canada. Canadian Journal of Zoology 55: 
628-630. 

Johnson, D., L. Kershaw, A. MacKinnon, and J. Pojar. 
1995. Plants of the western boreal forest and aspen park- 
land. Lone Pine Publishing and the Canadian Forest Service. 
Alberta, Canada. 

Kathman, R. D. 1990. Eutardigrada from Vancouver Island, 
British Columbia, Canada, including a description of Plati- 
crista chilensis n. sp. Canadian Journal of Zoology 68: 
1880-1895. 

McInnes, S. J. 1994. Zoogeographic distribution of terres- 
trial/freshwater tardigrades from current literature. Journal 
of Natural History 28: 257-352. 

Murray, J. 1910. Tardigrada of the British Antarctic Expedi- 
tion 1907-1909. London. 1: 81-185. 

Ramazzotti, G., and W. Maucci. 1983. The Phylum Tardi- 
grada. Memorie dell’ Instituto Italiano di Idrobiologia 41: 
1-1012. 

Richters, F. 1908. Beitrag zur kenntis der moosfauna Aus- 
traliens und der Inseln des Pazifishcen Ozeans. Zoologis- 
che Jahrbuecher Abteilung fuer Systematik Oekologie und 
Geographie der Tiere 26: 196-213. 

Van Rompu, E. A., W. H. de Smet, and J. M. Bafort. 1991. 
Contributions to the Tardigrada of the Canadian High Arc- 
tic. 2. Fresh water tardigrades from Little Cornwallis Island, 
Northwest Territories, Canada. Biologisch Jaarboek Dodon- 
aea 59: 132-140. 

Van Rompu, E. A., W. H. de Smet, and L. Beyens. 1992. 
Contributions to the Tardigrada of the Canadian High Arc- 
tic. 1. Freshwater Tardigrades from Devon Island, North- 
west Territories, Canada. Canadian Field-Naturalist 106: 
303-310. 

Whitehead, A. G., and J. R. Hemming. 1965. A comparison 
of some methods of extracting small vermiform nematodes 
from soil. Annals of Applied Biology 55: 25-38. 


Received 15 August 2004 
Accepted 14 November 2005 


*added in proof 2006 1105: 1-16 


2005 


NOTES 


589 


Mortality of Little Brown Bats, Myotis lucifugus, in a Rodent Trap in 


the Boreal Forest 


THomas S. JuNG! and BRIAN G. SLOUGH? 


'Yukon Department of Environment, Box 2703, Whitehorse, Yukon Y 1A 2C6 Canada; e-mail: thomas.jung @ gov.yk.ca 
35 Chronkite Road, Whitehorse, Yukon Y1A 5S9 Canada; e-mail: slough @northwestel.net 


Jung, Thomas S., and Brian G. Slough. 2005. Mortality of Little Brown Bats, Myotis lucifugus, in a rodent trap in the boreal 


forest. Canadian Field-Naturalist 119(4): 589-590. 


Accidental mortality of bats is not often observed or reported in the literature. It may, however, have an impact on population 
size and structure. We report an observation of 53 Little Brown Bats (Myotis lucifugus) being trapped and drowned in a home- 
made rodent trap at an abandoned cabin in southern Yukon. Traps of this design may be commonly used in the boreal forest. We 
recommend not using such traps in cabins that are used by colonies of bats. 


Key Words: Little Brown Bat, Myotis lucifugus, mortality, trap, Yukon. 


Accidental mortality of bats is not often observed 
or reported in the literature (Johnson 1933; Manville 
1963). Among the causes of mortality indirectly caused 
by humans (i.e. unintentional, or accidental mortality), 
several have been noted for bats. For example, poison- 
ing (e.g., Kunz et al. 1977; Clark et al. 1978; Pybus et 
al. 1986), human disturbance at roosting colonies or 
hibernacula (e.g. Tuttle 1979; Thomas 1995; Johnson 
et al. 1998), entanglement in exotic burdock (Arctium 
spp.; e.g., Lyon 1925; Hendricks et al. 2003), and colli- 
sions with wind turbines (e.g., Johnson et al. 2003), 
have been noted as observed or potential causes of 
accidental mortality. Curiously, few other sources of 
accidental mortality have been reported for bats. Yet, in 
some cases, accidental mortality may have an effect 
on colony size or structure that is equal to or greater 
than natural mortality (1.e., predation, disease, starva- 
tion, etc). Here, we report an observation of mortality 
of Little Brown Bats (Myotis lucifugus) in a homemade 
trap that is commonly used in buildings in the Canadi- 
an boreal forest to capture mice (Peromyscus spp.). 

On 8 September 2004 we found 53 dead and desic- 
cated Little Brown Bats in a rodent trap in a cabin near 
Squanga Lake, Yukon (60.5°N, 133.5°W). We were 
unable to reliably ascertain the age or sex of individu- 
als. The specimens have been retained by the Yukon 
Department of Environment. No rodents were found in 
the trap. 

The cabin was abandoned and housed a maternity 
colony of Little Brown Bats since at least 2000 (Slough 
2001*). Colony size before this incident of mortality 
was estimated at about 85-100 adults on 29 June 2004 
(Jung and Slough, unpublished data), based on emer- 
gence counts and captures with a harp trap (Tuttle 
1974). The rodent trap was prominently placed in an 
empty room that was regularly used for flight activity 
by bats after arousal and before they exited the cabin. 

The rodent trap was homemade and of a design that 
is commonly used in the Canadian boreal forest where 
remote cabins are left unoccupied for several months 
at a time, particularly by seasonal trappers, hunters, and 
fishers (H. Jessup, personal communication). These 
traps are regularly made of 19 litre (5 gallon) buckets 


FIGURE |. Homemade “bucket” trap with 53 mummified little 
brown bats (Myotis lucifugus). Note the wire and rotat- 
ing bait can that was used on the trap, but removed for 
this photograph. 


(31.4 cm in diameter by 38.7 cm tall) that are half- 
filled with water. A wire or rope is strung across the top 
opening of the bucket and a beverage can (approxi- 
mately 335 ml) is placed along the middle of the wire 
or rope. Rodents walk along the rope to reach bait that 
is placed in the can and fall into the bucket and drown. 
The trap is designed to catch multiple rodents, primarly 
Peromyscus. 

We suspect that Little Brown Bats attempted to 
drink from the bucket upon arousal from diurnal roost- 
ing and became trapped and drowned. Some bats, 
including Little Brown Bats, routinely drink after diur- 
nal roosting bouts and before foraging. Alternatively, 
some bats may not have attempted to enter the bucket 
to drink, but had tripped on the wire while flying low 
above the bucket and fell into the bucket and drowned. 
Trip-lines over water have been used as a live-capture 
technique for bats. Regardless, we believe that the water 
in the bucket was likely the attractant that led to bats 


590 


becoming trapped and drowning in the bucket. It is 
plausible that the distress calls of one bat trapped in 
the bucket attracted conspecifics, which then shared 
the same fate (Barclay 1982; B. Fenton, personal com- 
munication). Hendricks et al. (2003) suggested a sim- 
ilar scenario for multiple bats becoming entangled in 
burdock. 

Because Little Brown Bats tend to have long life 
spans and low annual productivity (Fenton and Barclay 
1980; van Zyll de Jong 1985), it is reasonable to sug- 
gest that accidental mortalities may, in some cases, 
have a substantial effect on colony size or structure. We 
suspect that this incident of accidental mortality may 
have substantially reduced the size of the Squanga Lake 
colony. We do not know how common it is to capture 
bats in this type of rodent trap, but suspect that it may 
happen somewhat regularly, with unknown effects on 
Little Brown Bat populations in remote boreal forest 
ecosystems. We strongly recommend that rodent traps 
of this or a similar design not be used in buildings 
when they are occupied by colonies of bats. 


Acknowledgments 

We thank B. Bennett for helping us solve the mystery 
of “how the bats got in the bucket”. Financial support 
was provided by the Yukon Department of Environ- 
ment and a Northern Research Endowment Grant from 
the Northern Research Institute, Yukon College. 


Documents Cited (marked * in the text) 

Slough, B. G. 2001. A survey of the bat fauna of the Yukon 
Territory: 2000 field studies. Unpublished report. Northern 
Research Institute, Yukon College, Whitehorse, Yukon. 
18 pages. 


Literature Cited 

Barclay, R. M. R. 1982. Interindividual use of echolocation 
calls: eavesdropping by bats. Behavoural Ecology and 
Sociobiology 10: 271-275. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Clark, D. R., R. K. La Val, and D. M. Swineford. 1978. 
Dieldren-induced mortality in an endangered species, the 
gray bat (Myotis grisescens). Science 199:1357-1359. 

Fenton, M. B., and R. M. R. Barclay. 1980. Myotis lucifugus. 
Mammalian Species 142: 1-8. 

Hendricks, P., J. Carlson, and C. Currier. 2003. Fatal entan- 
glement of western long-eared myotis in burdock. North- 
western Naturalist 84: 44-45. 

Johnson, P. B. 1933. Accidents to bats. Journal of Mammalogy 
14: 156-157. 

Johnson, G. D., W. P. Erickson, M. D. Strickland, M. F. 
Shepherd, D. A. Shepherd, and S. A. Sarappo. 2003. Mor- 
tality of bats at a large-scale wind power development at 
Buffalo Ridge, Minnesota. American Midland Naturalist 
150: 332-342. 

Johnson, S. A., V. Brack Jr., and R. E. Rolley. 1998. Over- 
winter weight loss of Indiana bats (Myotis sodalis) from 
hibernacula subject to human visitation. American Midland 
Naturalist 139: 255-261. 

Kunz, T. H., E. L. P. Anthony, and W. T. Rumage. 1977. 
Mortality of little brown bats following multiple pesticide 
applications. Journal of Wildlife Management 41: 476-483. 

Lyon, M. W. 1925. Bats caught by burdocks. Journal of Mam- 
malogy 6: 280. 

Manville, R. H. 1963. Accidental mortality in bats. Mammalia. 
27: 361-366 

Pybus, M. J., D. P. Hobson, and D. K. Onderka. 1986. Mass 
mortality of bats due to probable blue-green algal toxicity. 
Journal of Wildlife Diseases 22: 449-450. 

Thomas, D. W. 1995. Hibernating bats are sensitive to nontac- 
tile human disturbance. Journal of Mammalogy 76: 940-946. 

Tuttle, M. D. 1974. An improved trap for bats. Journal of 
Mammalogy 55: 475-477. 

Tuttle, M. D. 1979. Status, causes of decline and management 
of endangered gray bats. Journal of Wildlife Management 
43: 1-7. 

van Zyll de Jong, C. G. 1985. Handbook of Canadian mam- 
mals. Vol. 2: Bats. National Museum of Natural Sciences, 
Ottawa, Ontario. 212 pages. 


Received 18 March 2005 
Accepted 26 October 2005 


2005 NOTES 591 


Beavers, Castor canadensis, Feeding on Salmon Carcasses: 
Opportunistic Use of a Seasonally Superabundant Food Source 


JEFFREY S. GLEASON!, RYAN A. HOFFMAN’, and JAMES M. WENDLAND? 


‘Corresponding author: 9715 Independence Drive, Apartment B109, Anchorage, Alaska 99507 USA 
23738 D Gray Loop, Elmendorf AFB, Alaska 99506 USA 
3Alaska Department of Fish and Game, 333 Raspberry Road, Anchorage, Alaska 99518 USA 


Gleason, Jeffrey S., Ryan A. Hoffman, and James M. Wendland. 2005. Beavers, Castor canadensis, feeding on salmon carcasses: 
opportunistic use of a seasonally superabundant food source. Canadian Field-Naturalist 119(4): 591-593. 


We report observations of Beavers (Castor canadensis) foraging and feeding on discarded Chinook Salmon (Oncorhynchus 
tshawytscha) carcasses within the confines of the Susitna River drainage in southcentral Alaska on three separate occasions 
between 1999 and 2004. In all three instances, Beavers were observed actively seeking out freshly discarded carcasses or 
transporting “fresh” salmon carcasses in their mouths. In one instance, Beavers were seen using their dextrous forefeet to 
“handle” chunks of salmon while hunched over carcasses and in this case we actually witnessed Beavers “chewing” and 
ingestion was assumed. In the other two instances, Beavers were observed swimming with salmon carcasses in their mouths. 
Though unique within the framework of Beaver foraging ecology, we suggest this behavior may be a fairly common strategy 


employed by Beavers in Alaskan streams and rivers to take advantage of a seasonally superabundant source of protein. 


Key Words: Beaver, Castor canadensis, Chinook Salmon, feeding, Oncorhynchus tshawytscha, Alaska. 


Many herbivorous mammals are known to ingest 
animal matter either occasionally or on a seasonal basis, 
(i.e., mice [Peromyscus spp.; Barry 1977; Lackey et 
al. 1985], Red Squirrels [Tamiascirius hudsonicus; 
O’Donoghue 1994; Steele 1998; Yahner 2003] and 
Northern Flying Squirrels [Glaucomys sabrinus; Wells- 
Gosling and Heaney 1984], Muskrats [Ondatra zibethi- 
cus; Willner et al. 1980; Campbell and MacArthur 
1996; Erb and Perry, Jr. 2003], and White-tailed Deer 
[Odocoileus virginianus; Shaw 1963; Case and McCul- 
lough 1987; Pietz and Granfors 2000]). However, such 
observations are lacking for Beavers (Castor canaden- 
sis). The Beaver is considered a generalist herbivore, 
relying primarily on woody and herbaceous materials 
throughout its range to meet its energy requirements 
(Jenkins and Busher 1979; Novak 1987; Baker and Hill 
2003; Muller-Schwarze and Sun 2003). Unlike other 
herbivorous mammals that are known to ingest animal 
matter to meet seasonal food shortages or sex and age- 
specific nutrient requirements (Robbins 1993), obser- 
vations of carnivory are lacking for Beavers. We report 
here on observations of Beavers feeding on animal mat- 
ter in the form of Chinook Salmon (Oncorhynchus 
tshawytscha) carcasses and discuss the potential impor- 
tance of this readily available food source to Beavers 
in Alaska. 

On 5 July 2004, at around 0800 AST, we observed 
three Beavers feeding on Chinook Salmon carcasses 
discarded after being filleted by anglers along a rela- 
tively deep-water pool of Montana Creek located 
roughly 200 m upstream of the Parks Highway bridge 
(150°03'W, 62°06'N). The upper reaches of this creek 
and others in the Susitna River drainage represent 
important spawning areas for Chinook Salmon in the 
region. This stretch of the stream opened to salmon fish- 
ing on 3 July. Beavers fed on freshly discarded car- 
casses aground in shallow areas upstream and down- 


stream of the pool where the Chinook Salmon were 
holding. This pool was roughly 1.5 m at its deepest and 
was approximately 18 x 46 m in size, bounded on its 
upper and lower ends by shallow-water riffles. A vari- 
ety of both woody (willows [Salix spp.], birch [Betula 
spp.], alder [A/nus spp.]) and herbaceous vegetation 
[sedges (Carex spp.)] were abundant within the creek 
corridor. Weather conditions on this date were sunny 
to partly sunny with temperature ranging from 15 to 
20°C and no precipitation. The Beavers we observed 
appeared to be adults (based on their relatively large 
body size) and they fed for approximately 10-15 min 
on a carcass before moving on to another carcass. For- 
aging and travel between carcasses occurred nearly 
continuously for about 60 min in close proximity (5 m) 
to the observer (RAH). Chewing was evident as the 
Beavers fed while hunched over carcasses using their 
forefeet to “handle” chunks of remaining salmon mus- 
cle tissue (not organs). 

On two previous occasions approximately 6 km 
from the location of the above incident, we (JMW) 
observed apparent foraging behavior by Beavers that 
also involved Chinook Salmon. Two Beavers on 12 June 
1999 (07:00 ADT) and 2 on 17 June 2000 (04:00 ADT) 
swam from the Susitna River upstream into Goose 
Creek (150°05'W, 62°03'N) about 20 m to the edge of 
a 1.5 m deep pool. The creek was roughly 30 m wide 
and vegetation composition along the creek was similar 
to that previously described for Montana Creek. The 
Beavers, which all appeared to be adults, picked-up 
recently filleted Chinook Salmon carcasses discarded 
by anglers that were aground at the edge of the pool. 
The Beavers swam downstream from the pool into 
the Susitna River channel each with a carcass in its 
mouth. In addition, on 17 June 2000, we (JWM) ob- 
served a Beaver actually bite into an intact Chinook 
Salmon that was hanging from an angler’s stringer. The 


592 


angler and Beaver struggled briefly before the angler 
finally freed his catch. This bout resulted in two large 
(10-12 cm) gashes inflicted on the salmon from the 
Beaver’s incisors. 

We are unaware of other published observations of 
Beavers feeding on animal matter (see Jenkins and 
Busher 1979; Baker and Hill 2003). Given their mor- 
phological and physiological adaptations for feeding 
on woody and herbaceous vegetation (e.g., massive 
skull, dentition and large incisors, bacteria in cecum; 
Jenkins and Busher 1979; Novak 1987; Baker and Hill 
2003), this observation seemed significant in the con- 
text of Beaver foraging ecology. We assume that inges- 
tion of salmon muscle tissue occurred for at least one 
of the three observations since chewing was seen. We 
speculate that opportunistic feeding on salmon may not 
strictly be limited to Beavers in this watershed, but 
rather that it may be widespread in Alaskan rivers and 
streams where Beavers overlap large spawning runs of 
Pacific salmon. In fact, during our research we spoke 
with others (R. Prentki, Minerals Management Service) 
within the region who witnessed similar observations. 
We hypothesize that Beavers in Alaska and presum- 
ably elsewhere in the Pacific northwest may oppor- 
tunistically use or even actively seek out “fresh” (not 
senescent) salmon carcasses as a readily available and 
predictable (both in space and time) source of energy 
(i.e., protein or fat), at least on a seasonal basis (Gende 
et al. 2001; 2004b; Hilderbrand et al. 2004). Salmon 
carcasses, including discarded carcasses from anglers, 
represent a large, seasonal nutrient (particularly !°N) 
input into anadromous systems in Alaska and else- 
where (Cederholm et al. 1999; Hilderbrand et al. 1999; 
Helfield and Naiman 2001; Reimchen et al. 2003; 
Gende et al. 2004a). A diverse assemblage of birds and 
mammals utilize pre-spawning (predation) and post- 
spawning (scavenging) salmon for food (Willson and 
Halupka 1995; Hilderbrand et al. 2004). For example, 
Cederholm et al. (1989) documented forty-three taxa 
of birds and mammals on salmon streams in Washing- 
ton, and of these 51% were thought to have consumed 
salmon. Thus, it is not surprising that many animal 
species including apparently Beavers, exploit this sea- 
sonally superabundant high energy food source. 


Acknowledgments 

This manuscript benefited greatly from discussions 
with Howard Golden (Wildlife Biologist, Alaska 
Department of Fish and Game). H. Golden and K. F. 
Abraham (Wetlands Wildlife Scientist, Ontario Ministry 
of Natural Resources) provided comments on an earlier 
draft of this manuscript. Two anonymous reviewers 
provided suggestions that greatly improved the manu- 
script. 


Literature Cited 

Baker, B. W., and E. P. Hill. 2003. Beaver (Castor canaden- 
sis). Pages 288-310 in Wild mammals of North America: 
ecology, management, and conservation. Edited by G. A. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Feldhamer, B. C. Thompson and J. A. Chapman. Second 
Edition. John Hopkins University Press, Baltimore, Mary- 
land, USA. 

Barry, R. E., Jr. 1977. Length and absorptive surface area 
apportionment of segments of the hindgut for eight species 
of small mammals. Journal of Mammalogy 58: 419-420. 

Campbell, K. L., and R. A. MacArthur. 1996. Digestibility 
of animal tissue by muskrats. Journal of Mammalogy 77: 
755-760. 

Case, D. J., and D. R. McCullough. 1987. White-tailed deer 
forage on alewives. Journal of Mammalogy 68: 195-197. 

Cederholm, C. J., D. B. Houston, D. L. Cole, and W. J. Scar- 
lett. 1989. Fate of coho salmon (Onchorhynchus kisutch) 
carcasses in spawning streams. Canadian Journal of Fish- 
eries and Aquatic Sciences 46: 1347-1355. 

Cederholm, C. J., M. D. Kunze, T. Murota, and A. Sibatani. 
1999. Pacific salmon carcasses: essential contributions of 
nutrients and energy for aquatic and terrestrial ecosystems. 
Fisheries 24: 6-15. 

Erb, J., and H. R. Perry, Jr. 2003. Muskrats (Ondatra zibethi- 
cus and Neofiber alleni). Pages 311-348 in Wild mammals 
of North America: ecology, management, and conservation. 
Edited by G. A. Feldhamer, B. C. Thompson and J. A. Chap- 
man. Second Edition. John Hopkins University Press, Bal- 
timore, Maryland, USA. 

Gende, S. M., T. P. Quinn, and M. F. Willson. 2001. Con- 
sumption choice by bears feeding on salmon. Oecologia 
127; 372-382. 

Gende, T. P. Quinn, R. Hilborn, A. P. Hendry, and B. Dick- 
erson. 2004a. Brown bears selectively kill salmon with 
higher energy content but only in habitats that facilitate 
choice. Oikos 104: 518-528. 

Gende, S. M., T. P. Quinn, M. F. Willson, R. Heintz, and T. 
M. Scott. 2004b. Magnitude and fate of salmon-derived 
nutrients and energy in a coastal stream ecosystem. Journal 
of Freshwater Ecology 19: 149-160. 

Helfield, J. M., and R. J. Naiman. 2001. Effects of salmon- 
derived nitrogen on riparian forest growth and implications 
for stream habitat. Ecology 82: 2403-2409. 

Hilderbrand, G. V., S. D. Farley, C. C. Schwartz, and C. T. 
Robbins. 2004. Importance of salmon to wildlife: implica- 
tions for integrated management. Ursus 15: 1-9. 

Hilderbrand, G. V., T. A. Hanley, C. T. Robbins, and C. C. 
Schwartz. 1999. Role of brown bears (Ursus arctos) in the 
flow of marine nitrogen into a terrestrial ecosystem. Oecolo- 
gia 121: 546-550. 

Jenkins, S. H., and P. E. Busher. 1979. Castor canadensis. 
Mammalian Species 120: 1-8. 

Lackey, J. A., D. G. Huckaby, and B. G. Ormiston. 1985. 
Peromyscus leucopus. Mammalian Species 247: 1-10. 
Muller-Schwarze, D., and L. Sun. 2003. The beaver: natu- 
ral history of a wetland engineer. Cornell University Press, 

Ithaca, New York, USA. 

Novak, M. 1987. Beaver. Pages 282-313 in Wild furbearer 
management and conservation in North America. Edited 
by M. Novak, J. A. Baker, M. E. Obbard, and B. Malloch. 
Ashton-Potter, Concord, Ontario, Canada. 

O’ Donoghue, M. 1994. Early survival of juvenile snowshoe 
hares. Ecology 75: 1582-1592. 

Pietz, P. J., and D. A. Granfors. 2000. White-tailed deer 
(Odocoileus virginianus) predation on grassland songbird 
nestlings. American Midland Naturalist 144: 419-422. 

Reimchen, T. E., D. Mathewson, M. D. Hocking, J. Moran, 
and D. Harris. 2003. Isotopic evidence for enrichment of 
salmon-derived nutrients in vegetation, soil, and insects in 


2005 


riparian zones in coastal British Columbia. Pages 59-69 in 
Nutrients in salmonid ecosystems: sustaining production 
and biodiversity. Edited by J. Stockner, American Fisheries 
Society Symposium 34, Bethesda, Maryland, USA. 

Robbins, C. T. 1993. Wildlife feeding and nutrition. Academic 
Press, New York, New York, USA. 

Shaw, H. 1963. Insectivorous white-tailed deer. Journal of 
Mammalogy 44: 284. 

Shea, D. S. 1973. White-tailed deer eating salmon. Murrelet 
54: 23. 

Steele, M. A. 1998. Tamiasciurus hudsonicus. Mammalian 
Species 586: 1-9. 

Wells-Gosling, N., and L. R. Heaney. 1984. Glaucomys sabri- 
nus. Mammalian Species 229: 1-8. 


NOTES 


393 


Willson, M. F., and K. C. Halupka. 1995. Anadromous fish 
as keystone species in vertebrate communities. Conserva- 
tion Biology 9: 489-497. 

Willner, G. R., G. A. Feldhamer, E. E. Zucker, and J. A. 
Chapman. 1980. Ondatra zibethicus. Mammalian Species 
141: 1-8. 

Yahner, R. H. 2003. Pine squirrels (Jamiasciurus hudsonicus 
and 7: douglasii). Pages 268-275 in Wild mammals of North 
America: ecology, management, and conservation. Edited 
by G. A. Feldhamer, B. C. Thompson and J. A. Chapman. 
Second Edition. John Hopkins University Press, Baltimore, 
Maryland, USA. 


Received 4 November 2004 
Accepted 5 December 2005 


Book Reviews 


ZOOLOGY 


Arrivals and Rivals: A Birding Oddity 


By Adrian Riley. 2005. Brambleby Books, PO Box 410, 
Harpenden, Hertfordshire, ALS SXQ UK. 165 pages. £7.99 


paper. 


Many of you will know I am not a lister (or twitcher 
as they are called in Britain.). | am nowhere near com- 
petitive enough. I really do not care if you have seen 
more than me, as long as I see what I want to see. 
This is a book about the author’s efforts to meet the 
ultimate twitcher’s challenge — to become the British 
Twitcher of the Year. So I was intrigued as to why he 
did it and against what social backdrop. 

The author starts his year with his retirement and a 
consequent move to Norfolk. Now Norfolk is a very 
good location for a birdwatcher as it has an interesting 
list of annual birds and gets a great number of vagrants. 
The author starts his year as a very keen birder who 
keeps an annual list. His early partnership with an avid 
twitcher nets him a very respectable winter list by the 
end of February. His retirement allows him the time to 
chase many rarities, particularly those on his new back 
door step. 

Then his tone starts to change as he become more 
greedy for ticks. He pushes himself harder and farther 
to add a new bird. He finds he has become a “combat 
“birder, where the tick transcends everything else. Friend- 
ships pull apart and then become bitter and paranoid. 
He becomes over-emotional and over-tired. The com- 
petition becomes a war and many of the joys of bird- 
ing fall to the wayside. The end of December comes 
as a relief. The fighting is over and he has won with a 
tally of 380 compared to the 573 on the official list of 
birds of Great Britain sanctioned by British Ornithol- 
ogists’ Union’s Records Committee (BOURC). 

His total is over two-thirds of the British list. Or is it? 
He includes one bird that had not made it to the official 
list by November 2005 (Elegant Tern, although this 
California to Chile bird is on the Irish list — and it 
appears “Britain” still includes Ireland for ticklisters. 
The BOURC however states “Species recorded from 
the Republic of Ireland (jurisdiction of the Irish Rare 
Birds Committee (IRBC)) and the Channel Islands are 
not covered in any reports or lists produced by the 
BOURC). There are two species (Baikal Teal and 
White-headed Duck) for which there is “reasonable 
doubt” and therefore “they form no part of the British 
list” (quotation marks are BOURC text). He has also 
added five species whose taxonomy as a full species is 


still in debate. Again the BOURC has not seen fit to add 
them at this ttme. BOURC has split redpoll into three 
species, but most authorities, including the American 
Ornithologists Union and Birdlife International, recog- 
nize only two species saying the taxonomy of the Less- 
er Redpoll is no longer valid. His total is submitted for 
scrutiny, but I am not clear who does this or what ref- 
erence they used to make his list “official.” 

I was amazed at how well-organized rarity chasing 
is in Britain. Although the author does not explain the 
system, I deduced there is a manned Rare Bird Alert 
(RBA) that accepts and vets calls, issues bulletins by 
pager and keeps the information up to date. This means 
a birder chasing a vagrant in the far north can receive 
an alert with instructions and drive to the south coast 
for another tick on the same day! 

The cost of this epic is that he has seen the darkest 
side of himself and others. Withholding or obscuring 
RBA information is only part of the “game.” He en- 
dured a great deal of discomfort, potential legal action, 
sloshing through that awful British weather and in driv- 
ing many long, lonely hours. As well it cost him friend- 
ship and about $16 000 Canadian. 

To his credit, he maintained a sense of humour and 
never really lost the true meaning of birding (although 
you had to wonder on occasions!). He also never low- 
ered himself to questionable sightings or inclusions. 
Questionable practices is a different question. 

I was amazed at the number of vagrants that turned 
up in Britain in a single year. Over 30% of his ticks were 
single sightings and another 11% were two sightings 
only. 

I was truly fascinated by this book. It is certainly a 
book for all birders — sane and insane — and definitely 
a must for all marathon twitchers. I enjoyed the writer’s 
style and I hope he takes on a new project with a dif- 
ferent, more valid objective that leads to another book 
(perhaps he could retrace the steps of Henry Seebohm; 
Siberia in Asia: A Visit to the Valley of the Yesnay in 
East Siberia, with Description of the Natural History, 
Migration of Birds, etc. 1882?). Then he would avoid 
adding his last sentence in the book — “In the end, it 
matters not a jot.” 

Roy JOHN 
2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 
K1J 6K5 Canada 


594 


2005 


Band-tailed Pigeon: Wilderness Bird at Risk 


By Worth Mathewson. Timber Press, Portland, Oregon. 2005. 
Hardcover. 183 pages, 20 colour photographs, 23 black- 
and-white figures, 4 tables. $19.95 US. 


Ward Mathewson has been studying the Band-tailed 
Pigeon for 48 years, first as one who hunted them and 
enjoyed eating pigeon pot-pie, and later as a conserva- 
tionist trying to save the species. We learn much about 
the year-round habits and habitat of this beautiful bird. 
It breeds north to extreme southwestern British Colum- 
bia, but occasional individuals wander farther north and 
east, into the three prairie provinces. Mathewson has 
searched the literature carefully, and cites three Sas- 
katchewan bandtail records in the Blue Jay. In keeping 
with other dove species, the bandtail builds a flimsy nest 
and the female generally lays a single egg. Favourite 
foods are berries of the cascara and elder; because of the 
extremely high potassium content of those berries, band- 
tails have a physiologic need to visit mineral springs with 
a compensatory high sodium (and calcium) content. 

Initially, in the 1890s bandtails were subjected to net- 
ting and shooting over decoys for distant markets. As 
late as 1912, a train would bring 100 hunters to a choice 
habitat and they would shoot an average of 30 birds 
each. One hunter shipped 2000 bandtails to hotels in 
California during the winter of 1911-1912. Until farm- 
ers learned to drill seeds into the ground, this pigeon 
was persecuted for eating a high proportion of hand- 
sown seeds spread on the ground. Later, bandtails des- 
troyed up to nine-tenths of cherries in a grove until 
the farmer invited 500 hunters to shoot the bandtails. 
In subsequent years, sport hunters often concentrated 
their efforts at mineral springs and in mountain passes 
through which bandtails migrated. 

There were early warnings about declining numbers. 
Johnson Neff in California in 1932 and 1934 warned 
that as few as one in five bandtails mortally shot were 
actually retrieved by the hunter. Art Einarsen in 1946 
reported than at one prime 10-acre bandtail hunting site 
near Crawfordsville, Oregon, 10 000 shells had been 
fired at a 10-acre shooting site. Gene Silovsky deter- 
mined in 1969 that 52 of 74 males and 48 of 82 females 
shot in the fall hunting season were actively produc- 
ing pigeon milk, hence were feeding squabs; young 
inevitably died if no parent returned to feed them. 
Government game departments simply ignored such 
research results. 

In 1975, when the new Oregon Fish and Wildlife 
Commission was formed, author Worth Mathewson 
began his campaign to have shooting stopped at the few 


BOOK REVIEWS 


395 


mineral springs where bandtails concentrated. He felt 
shooting here during the first two weeks of September 
led to unsporting slaughter, certainly not “fair chase,” 
as waves of males and then females fell to the con- 
centrated hunters’ guns. As numbers declined, the daily 
bag limit was lowered from eight to five birds per day, 
without apparent effect. In 1980 and 1981, the open- 
ing day was delayed two weeks to 15 September, to a 
time when fewer squabs were being fed; there was a 
noticeable increase in the population index. But self- 
interest of the bandtail shooters persuaded the state to 
change the opening date back to September | for eight 
more destructive years. In 1989, the 15 September 
opening was restored, the season length was shortened 
to seven days and the bag limit reduced to two birds, 
yet despite these changes a federal Fish and Wildlife 
Service biologist observed that “We’re going to ‘do- 
nothing them’ into extinction.” By 1991, numbers had 
dropped so much that British Columbia and Washing- 
ton closed their seasons. The bandtail had been a vic- 
tim of stupidity, ignorance, misrepresentation, and in- 
attention to research findings. 

Meanwhile, forest companies which wanted a high- 
er harvest of conifers, began large-scale aerial spraying 
of broad-leaved trees in the 1970s. The Nehalem Val- 
ley, a bandtail stronghold, became solid conifers, with 
the elderberry and cascara apparently gone forever. 

This book is much more than the frustrating crusade 
by Mathewson on behalf of the Band-tailed Pigeon. 
It is also an example of investigative journalism at its 
best, well-documented and convincing. Mathewson 
shares with us the inside stories, and compares the 
bandtail situation with that of other doves and pigeons, 
including the extinct Passenger Pigeon, and the de- 
creasing numbers of the White-crowned Pigeon, White- 
winged Dove, and Key West Quail-Dove. Meanwhile, 
in Argentina 210 shooters tallied 408 643 Eared Doves 
in one season, described by Mathewson as “mindless 
killing without justification.” As late as 2003, an issue 
of Gray’s Sporting Journal carried an advertisement 
picturing a shooter kneeling on 7696 Eared Dove car- 
casses! 

The writing is both lyrical and descriptive, a pleasure 
to read. Sketches by David Hagerbaumer add to the 
attractiveness of this book. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N OJ8 
Canada 


596 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Monitoring Bird Populations Using Mist Nets: Studies in Avian Biology #29 


Edited by C. John Ralph and Erica H. Dunn. 2004. Cooper 
Ornithological Society, c/o western Foundation of Verte- 
brate Zoology, 439 Calle San Pablo, Camarillo, Califor- 
nia 93012-8506 USA. 211 pages. U.S.$39.95. 


A workshop was held in California in October 1993. 
Forty experts (six from Europe, five from Canada, 1n- 
cluding one of the two editors, and the remainder from 
the United States and Puerto Rico) evaluated the strength 
and weaknesses of mist-netting as a method of moni- 
toring bird populations. Six additional individuals con- 
tributed to the papers, incorporating new data and up- 
dated analyses prior to final editing — which occurred 
only after a remarkable delay of ten years. 

As stated by the editors in their Introduction, the 
advantages of mist nets include ease of standardized 
sampling, low observer bias, ability to detect species 
that are often missed (sometimes including rarities un- 
likely to have been spotted by routine observation), 
and the opportunity to examine birds in the hand. Mist 
net studies, as do other visual and aural methods, pro- 
vide indices of abundance rather than total counts of 
populations. 

Three programs with some variation in methodolo- 
gies, but using mist-nets in a standardized fashion, 


are the MAPS (Monitoring Avian Productivity and 
Survivorship) Program pioneered in California by 
David DeSante and now spreading across the continent, 
the British Trust for Ornithology’s Constant Effort 
Sites Program, and the MRI Program in continental 
Europe (named from the first letter of the three initial 
sites, since expanded to seven). All three methods have 
the potential to detect long-term temporal trends. Sam- 
pling only at weekends should be avoided. 

Two major papers deal with 18 years of data col- 
lected at Long Point Bird Observatory in Ontario. At 
Beaverhill Bird Observatory in Alberta, six participants 
in an informal banding training program failed to attain 
fully satisfactory performances. Summary recommen- 
dations, to produce as much information as possible 
with the lowest bird mortality, include training of all 
participants, checking the nets every 15 to 30 minutes, 
and use of 30- and 36-mm-mesh nets that are 12 m long. 

This volume is a must for anyone using mist nets, 
and for Bird Observatory libraries. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


Portraits of the Bison: An Illustrated Guide to Bison Society 


By Wes Olson. 2005. The University of Alberta Press, 
Edmonton. 108 pages. ISBN 0-88864-432-9. Paperback. 
$39.95. 


Bison are the iconic animal of the Canadian prairies. 
If you live in western Canada, then pictures of bison 
— being hunted, as piles of skulls by train tracks, as a 
provincial symbol — are woven into your conscious- 
ness. Yet, most people know little about them. It is this 
deficiency that Wes Olson has set out to remedy with 
this full-colour guide. Olson has worked as a park war- 
den at Elk Island National Park, located just east of 
Edmonton, for many years. Despite its name, the Park 
is probably best known as the home of two bison herds, 
one of plains bison and the other of wood bison. 
Olson’s experience in working with these herds has 
given him unique insight into the social structure and 
biology of the animals. At the same time, his interac- 
tions with park visitors have made him aware, as he 
explains in the introduction, that most people do not 
treat bison with the respect they deserve and thus some- 
times get themselves into dangerous situations. These 
are the two incentives that encouraged him to com- 
pile this volume. Wes Olsen is a talented artist and the 
book is abundantly illustrated with his drawings of 
bison, providing vivid evidence of keen observation 
and long study. His wife, Johane Janelle, contributed 
many outstanding colour photographs, capturing bison 
in different moods and surroundings. 

The book is arranged in four chapters. Olson pro- 
vides a short introduction to bison and notes the occur- 


rence and ranges of the two main modern forms, plains 
and wood bison. Interestingly, he describes the capture 
and establishment of the wood bison herd, but does 
not go into the history of the plains bison recovery in 
any detail. I found this omission curious, given that the 
rest of the volume concentrates on plains bison, espe- 
cially since saving the bison from the brink of extinc- 
tion is a classic conservation story. The second chapter 
focuses on safety and awareness for hikers and others 
who might run into bison in the field. Olson describes 
the stages of a bison’s reaction to an encounter with a 
human. He emphasizes that bison have a very large 
“personal space” and should be given an extremely 
wide berth. Olson identifies three zones within “bison 
space” — the awareness zone, the escape zone, and the 
fight zone — and indicates what a bison’s behaviour 
may be when a human impinges on each of these zones. 

The next two chapters comprise the heart of the 
book. The first is focused on the seasonal cycle and 
structure of bison herds. This makes it clear that bison 
have a complex social structure, related to biological 
events, such as calving and the rut, and the life-stages 
of the animals. The next chapter provides more detail 
on the life-stages of a male and a female plains bison. 
This is the longest chapter (34 pages) and 1s richly 
illustrated with page by page pictures of bison, usually 
acutely-detailed profile drawings with the salient iden- 
tifying features indicated. Olson comments that dis- 
tinguishing male and female bison is not as straight- 
forward as one might think, especially when the bison 


2005 


are seen at a distance or in a large group. For exam- 
ple, although males are usually larger than females, 
this size difference is not as marked in younger ani- 
mals. Olson distinguishes seven life-stages for bison: 
calves, yearlings, two-year-olds, young adults, mature 
adults, dominant males, and aged bison. He points out 
that for wildlife biologists, knowing the age structure 
and composition of a bison herd is important for mak- 
ing management decisions. However, recognizing dif- 
ferent generations of bison can also add interest to a 
hike or nature ramble. Just as birders recognize birds 
by plumage and songs, bison watchers must learn to 
distinguish different pelage stages, body structure, and 
horn shapes. 

The book continues with four appendices, focusing 
on age and sex determination from the skull, a list of 
public bison herds in North America, a summary re- 
iterating the main features of male and female life 
stages, and a blank data sheet for recording observa- 
tions on bison herd structure. The age and sex deter- 
mination focuses on a few critical plains bison skull 
measurements and tooth eruption and wear patterns 
in the lower jaw or mandible. Olson mentions one cri- 
terion for distinguishing bison from cattle skulls. I 
found this section a bit brief. Within a national park 
or a wildlife refuge with a bison herd one can assume 
that skulls are mostly going to be from bison. How- 
ever, this is not the case across the prairies at large, 
where people often find skulls, or more usually parts 
of skulls, that they think may be bison. I felt that more 
information on distinguishing bison from other faunal 
remains would have made this section more generally 


BOOK REVIEWS 


398 


useful. The volume concludes with a helpful reference 
list, comprising many general and scholarly works on 
bison behaviour and biology. The book is large-for- 
mat (27.5 cm wide by 25.5 cm high), giving plenty of 
space for the photographs and drawings, and is print- 
ed on high-quality glossy paper. The images are well 
laid-out and, for the most part, crisply rendered. 

This is an unusual book. I certainly have not come 
across anything quite like it before. It is not really a field 
guide, since the format does not lend itself well to being 
stuffed in a pocket. It is not really a text book, because 
the emphasis is on identification, observation, and field 
work. This book will certainly be relevant to anyone 
who works with bison in the field or has an interest in 
bison population studies, wildlife biology, or ecolo- 
gy. However, I think it has much broader appeal and 
will also intrigue anyone who simply likes watching 
bison and wants to learn more about them. The book 
can also be enjoyed on a purely aesthetic level, because 
the photographs and drawings are in themselves so 
attractive and fascinating. Those of us fortunate enough 
to live near Elk Island National Park can see bison in 
the flesh, large lumbering presences moving through 
aspen groves, or dozing in summer sunlight. Armed 
with information from this book, bison spotting can 
have another dimension, as you try to work out who’s 
who in the herd. 


ALWYNNE B. BEAUDOIN 


Royal Alberta Museum, 12845-102"' Avenue, Edmonton, 
Alberta TSN OM6 Canada 


The Buffalo Wolf, Predators, Prey and the Politics of Nature 


By Lu Carbyn. 2003. Smithsonian Books, Washington, D.C. 
USA. 248 pages. U.S.$24.95. Cloth. 


“It might be said that the wolf was one of the last 
natural resources to be included in the great modern 
movement toward conservation.” This statement ap- 
peared in the late 1960s in one of the most impressive 
and enjoyable books about wolves that has ever been 
written (Rutter and Pimlott 1968). Conservation has 
come a long way, both generally and specifically with 
regard to the wolf. People who have led the way are 
few in number. Lu Carbyn is one of them. His devo- 
tion to conservation and wilderness is evident in this 
book which centres on wolves and bison in the Peace- 
Athabasca delta region of Wood Buffalo National Park. 

The first chapter contains some very graphic des- 
criptions of wolves attacking and killing bison. This 
is disturbing, but the reader is challenged to accept 
that the health of the ecosystem lies in the survival of 
the predators. In contrast, the second chapter sets the 
stage, describing Wood Buffalo Park and providing 
some biological background. The real story begins 
with the third chapter. It presents observations made 
during the early days of the research in Wood Buffalo 


Park. The social structure of the pack as it relates to 
the interactions with bison is described in some depth. 
The fourth chapter outlines some of the fascinating 
results of aerial surveys. Chapters 5, 6, 7 and 8 mostly 
have to do with field studies that followed the termi- 
nation of the Canadian Wildlife Service program in 
1984. Adventures with film crews are a major feature. 

Chapter 9 includes some experiences during later 
solo visits to the delta. The debate over whether or 
not to eradicate the Wood Buffalo bison herd is the 
main focus of the last three chapters. This debate had 
its origin in the introduction and “contamination” of 
the park’s wood bison with plains bison stock from 
Wainright. Not only were the park bison impure, they 
carried diseases that also infect cattle. On the other 
hand there may always have been hybrids and maybe 
it is unlikely that herds can be protected indefinitely 
from some or all diseases so better to let nature take 
its course. Alleged declining numbers of bison in the 
park fanned the fire of eradication, but “how much 
management is too much?” After years of first hand 
study Carbyn believed that the wolves were the main 
factor in the fluctuating numbers of bison but he point- 
ed out that parts of the big picture are still not avail- 


598 


able and more information is needed to predict the 
future of the park bison. The author cleverly leads us 
through this debate showing how views changed and 
how personal agendas and politics have played a role 
at the expense of science. He concludes that “if other 
human priorities are set in place, then enlightenment, 
not agendas, should rule the day.” Selected references 
and an index help to make this book a source of en- 
lightenment for the management of the buffalo wolf 
ecosystem. 

The book will appeal to a wide audience and is 
perhaps structured to do that. As a biologist I would 
have appreciated a little more development in some 
areas, perhaps at the expense of information about 
people. The differentiation of Plains and Wood bison 
was not sufficiently explained. There are some inter- 
esting aspects to discuss here (for more information 
on separating the two kinds of bison and other infor- 
mation, see Mitchell and Gates 2002). As another ex- 
ample, much more could have been said about the 
starlings observed perching on the bison in chapter 3. 
They are a reminder of the impact of humans, even in 
this remote wilderness. They spread from a flock intro- 
duced to New York City in 1890. At the time this may 
have been the furthest north that they had spread in 
Canada and clearly they were not confined to human 
habitations such as Fort Smith. This is additionally 
interesting because it is a reminder of another care- 
less introduction which contributed later to the major 
bison controversy in Wood Buffalo Park. Although I 
would like to have seen more in some areas, the book 
is not without anecdotes including subjects such as 
pollination of waterlilies, history of bush pilots, change 
in traditional experience of nature by native people 
and movement distances of lynx. etc. It is probably a 
good thing that Carbyn is able to focus to the extent 
that he does because there is so much to say in the 
experience of nature in the Canadian north. 

Like most dedicated biologists and teachers, Carbyn 
has taken a number of hard hits. One of these was 
when the CWS wolf study program was terminated. 


Grassland Grouse and their Conservation 


By Paul A. Johnsgard. 2002. Smithsonian Institution Press, 
Washington, D.C. Hardcover. 157 pages. $34.95 U.S. 


When Paul Johnsgard perceived that most grassland 
races of grouse had decreased in numbers since he 
wrote The Grouse of the World in 1983, and that some 
were apparently doomed to follow the “Heath Hen” 
into extinction, he responded by writing yet another 
book. Johnsgard is well qualified for the task. He 
writes lyrically, draws exquisite sketches, and is cited 
in the Seventh American Ornithologists’ Union Check- 
List as an authority on the taxonomy of the Greater 
and Lesser prairie-chicken. 

The first chapter deals with the decline of the 
Heath Hen and its final extirpation on Martha’s Vine- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Of course many government programs are not based 
on long term visionary thinking, but I am not sure 
that I can accept Carbyn’s view that this is usually a 
consequence of leaders lacking vision. Leaders are 
often just not strong enough to break away from the 
constraints of ignorance surrounding them. Without 
actually saying so, Lu Carbyn reminds us that inde- 
pendent and visionary thought as well as implementa- 
tion of personal goals are characteristic of some peo- 
ple. Fortunately, these people help to compensate for 
the lack of long term visionary thinking elsewhere. 
Carbyn has a message for biologists. He describes how 
in field study, intuition and imagination play an impor- 
tant role. We are reminded to think like a scientist — 
but not to the exclusion of being a naturalist. He notes 
that “we should be taking from nature... a sense of 
what it is rather than its soul.” The style of writing is 
personal and conversational with digressions and flash- 
backs. It is a very pleasant read. 

This book promotes wilderness which is a part of 
Canadian heritage and identity and this alone makes 
it a valuable contribution. It also presents a fascinat- 
ing biological and historical perspective on wolves and 
bison in Wood Buffalo Park. It introduces the world 
of a biologist. It will bring to many readers an under- 
standing of the science, issues and politics of biology. 
Anyone can learn from it and enjoy it. By educating 
in numerous ways it will contribute to the protection 
of nature. It should be a source of pride for Canadian 
field biologists. 


PAuL M. CATLING 


170 Sanford Avenue, Ottawa, Ontario K2C OE9 Canada 


Literature Cited 

Mitchell, J. A., and C. C. Gates. 2002. Status of the Wood Bison 
(Bison bison athabascae) in Alberta. Alberta Sustainable Resource 
Development, Fish and Wildlife Division, and Alberta Conserva- 
tion Association, Wildlife Status Report Number 39, Edmonton, 
Alberta. 32 pages. http://www3.gov.ab.ca/srd/fw/status/reports/bison/ 
body.html 

Rutter, R. J., and D. H. Pimlott. 1968. The world of the wolf. J. B. 
Lippincott Company. New York. 202 pages. 


yard, off the coast of Massachusetts. The remainder of 
the book provides us with a detailed account of recent 
population trends among the various races of grassland 
grouse, and provides current rough population estimates 
for each. Such detailed state-by-state accounts of de- 
clining ranges and populations for almost all races 
makes for depressing reading. 

The Lesser Prairie-Chicken, whose closest ecological 
partners are Shinnery Oak and Sand Sage, is holding 
its own in one part of Kansas, but its range has de- 
creased greatly in each of its five states. In New Mex- 
ico, the population has dropped from 40 000 to 1000; 
in Oklahoma, from 15 000 to less than 3000; in Col- 
orado, numbers have dropped by about 97 per cent; 
in Texas it has dropped to about one-third of one per 


2005 


cent of its historic numbers. In a few areas of contact 
with its larger relative, the Greater Prairie-Chicken, 
some hybridization occurs. 

The Greater Prairie-Chicken vanished from Ten- 
nessee by 1850, Kentucky by 1874, Arkansas by 1913, 
Ohio by 1934, and Michigan by 1983. Hunting sea- 
sons were closed permanently in Missouri in 1907, 
Colorado in 1937, Minnesota in 1943, North Dakota 
in 1946, Wisconsin in 1956, and Oklahoma in 1998. 
The last known nesting in Iowa was in 1952, but after 
reintroduction attempts there are now 44 males on six 
booming grounds. In Kansas, Nebraska and South 
Dakota, populations remain large enough to permit an 
annual hunting season, but even in these three states 
the area occupied has decreased drastically. 

The attwateri subspecies once may have numbered 
100 000 birds when it occupied six million acres of 
Gulf Coast prairies, mainly in southeastern Texas. Now 
that predators take 53 per cent of nests, brood survival 
to eight weeks is 34 per cent, and adult annual survival 
rates vary from 11 to 36 per cent, the population has 
dropped to about 45 birds. This race appears to be 
doomed. 

The Greater Sage-Grouse, specifically adapted to 
sagebrush, has dropped from a population of near 2 
million in 1950 to about 200 000 in 2000, and it has 
been extirpated from Nebraska, Oklahoma, New Mex- 
ico, and Arizona. In the late 1970s it was still being shot 
by hunters at the rate of 83 O00 per year in Wyoming 
alone. In Saskatchewan, the population dropped by up 
to 92 per cent between 1970 and 2000. 

In 2000, when the Gunnison Sage-Grouse was 
named a new species, Centrocercus minimus, it occu- 
pied only a small portion of its former range in south- 
west Colorado and adjacent Utah. Fewer than 5000 
adults remained. Johnsgard notes that this new species, 
due to the “glacial rate of movement of the federal 
legal process ... might well be endangered before [it 
is] officially recognized as threatened, and extinct 
before [it is] classified as endangered.” 


Crows: Encounters with the Wise Guys 


By Candace Savage. 2005. Greystone Books #201-2323 Que- 
bec Street, Vancouver, British Columbia VST 4S7 Canada. 
113 pages. $27 Cloth. 


Several years ago I overheard a person say “He must 
have been slow to volunteer. He’s got stuck with 
crows.” They were talking about me. I had actually 
been one of the first to volunteer to write sections of 
the original Ontario breeding bird atlas and I had 
deliberately asked for crows as my assignment. Why 
do people think of these birds as common, black and 
noisy, and therefore dull? 

I have always found crows to be entertaining, clever 
and well worth watching. It is comforting to know there 


BOOK REVIEWS 


a9 


Finally, two of the three grassland subspecies of 
the Sharp-tailed Grouse have shown drastic decreas- 
es in range, including a loss of 90% of the range of 7: 
p. campestris in Wisconsin and Michigan and 60% in 
Minnesota, even though their habitat has changed less 
than that of any other grassland grouse species. In the 
far west, an even more drastic loss of range (92%) has 
occurred for 7. p. columbianus. Since 1998 it has been 
listed as threatened in the state of Washington; satis- 
factory populations survive only in British Columbia. 

The greatest strength of this book is the detailed 
maps of past and current ranges of each race. Descrip- 
tions of the mating rituals of each race are especially 
well done. I have minor criticisms. Coverage is less 
detailed for all grassland grouse in the Canadian 
provinces; Johnsgard fails to state conclusively that 
the Greater Prairie-Chicken is officially considered to 
be extirpated from Canada, and that the recovery 
team for this species was disbanded in 1994. It is not 
true that hybridization between Greater Sage-Grouse 
and Sharp-tailed Grouse occurs only in eastern Mon- 
tana and eastern Wyoming; it has also been reported 
from extreme southern Saskatchewan. The disruptive 
effect of Ring-necked Pheasant cocks at Greater Prairie- 
Chicken leks is barely mentioned. 

The final chapter asks, “Can the fabric be mended?” 
For some races of grassland grouse, the Greater Sage- 
Grouse in Canada, the Lesser Prairie-Chicken and the 
attwateri race of the Greater Prairie-Chicken in the 
United States, it may already be too late. “As agriculture 
became more mechanized and the farms larger, fewer 
and fewer acres were spared from increasingly effec- 
tive pesticides and herbicides ... the prairie grouse 
began a long, slow, but certain descent into oblivion.” 

No one knows what the future holds, but Johnsgard 
certainly has raised the alarm call. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N OJ8 
Canada 


is at least one person who shares my enthusiasm for 
these wonderful birds. Candace Savage has collected 
tales and myths about the crows of our world. For the 
crow fan this is delightful reading. For the crow-defi- 
cient minds of others this book will offer enlightenment. 

The author has sifted though both archaic stories 
and modern research on the black crows (the gaudy 
jays do not get mentioned). She covers the biology, 
social structure and interactions, communications and 
tool-use to try and answer the key question. Are crows 
intelligent? Crows have been observed by many of us 
doing things that suggest they have mentally resolved 
a problem. The difficulty is their behaviour can be 
explained in other ways. The research that Savage 


600 


references is generally aimed at giving a crow a test 
that should prove or disprove intelligence. This is an 
elusive goal and I am not convinced that crow intelli- 
gence is proven. However, the results are so amazing 
that it is impossible to deny that crows behave intelli- 
gently. In some ways they are surprisingly similar to 
humans. 


Antipredator Defenses in Birds and Mammals 


By Tim Caro. 2005. The University of Chicago Press, Chicago 
60637. xv + 591 pages. U.S. $95.00 Cloth. $38.00 Paper. 


Recently I read a report of Merlins hunting migrant 
chickadees as they flew out from a point along Lake 
Ontario. The falcons were capturing and eating the 
chickadees on the wing, and it made me ponder how 
defenseless the birds seemed, and the contrast with the 
shorebirds that Merlins are normally hunting in this 
location. To those of us who watch birds such events 
can often seem exciting and dramatic interludes in 
the birds’ more usual activities, such as feeding. As 
such, they have generated over the years a host of anec- 
dotal notes of the type above. But of course, defenses 
against predators are absolutely basic to an animal’s 
life: without them there would be no feeding, or any- 
thing else. So the challenges of avoiding predation 
have become a fertile field of investigation. 

Which brings me, finally, to this book, in which Tim 
Caro examines the bewildering range of mechanisms 
and strategies that enable mammals and birds to evade 
capture and death from the wide variety of predators 
that most animals face. Such is the multiplicity of all 
of these — prey species, defenses and predators — that 
any synthesis is daunting, yet Caro does a masterly job 
of mustering the relevant data from a huge and grow- 
ing field, and rendering it into a coherent and lucid 
whole. The 87 pages of references alone attest to the 
encyclopedic character of the undertaking. 

The book is divided into 13 chapters, with the first 
devoted largely to setting the stage, followed by 11 that 
trace the “predatory sequence” from traits that avoid 
detection, such as cryptic coloration, and behavioral 
mechanisms to avoid detection, through vigilance and 
prey signaling [both warning and of unprofitability], 
the benefits of grouping, and then physiological and 
morphological defenses, such as body size, quills, 
spines, antlers and the like. There is a chapter on nest 
defense, and one on mobbing and other group defens- 
es, with chapter 12 devoted to “behaviors of last resort’. 
Taken together, these provide a comprehensive over- 
view of predator avoidance, with copious references 
to the literature, and tables and figures from relevant 
papers providing a useful complement to the text. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


This is an ideal gift book. It is fun to get and read, 
but not the sort of book you normally buy for yourself. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 
K1J 6K5 Canada 


Caro’s formidable command of his subject is appar- 
ent throughout. 

The thing that impressed me most about the book 
is its clarity. As a generalist with no more than a broad 
interest in the field, I had some trepidation about un- 
dertaking this review. The subject is, after all, at this 
level a technical one, and technical treatises can be 
relatively incomprehensible to the non-specialist. I need 
not have worried. While hardly light bedside reading, 
the entire text has a refreshing clarity and readability. 
From the preface onwards Caro clearly states what 
he is going to do, with the reasons why, and provides 
the appropriate references to those allied aspects of 
the field that he chooses not to cover. Then in each 
chapter he again defines the area he will cover, dis- 
cusses it critically with copious examples, noting the 
weaknesses in our current understandings, and then 
provides a concise summary at the end. 

Chapter 13, framing questions about antipredator 
defenses, brings the disparate threads of the previous 
chapters together, and focuses in on ten “pressing ques- 
tions” which the author sees as particularly important 
at this time. 

An appendix gives the scientific names of verte- 
brates mentioned, and each chapter has a delightful 
sketch of some aspect of animal behaviour that is 
appropriate to the text that follows. All the references 
that I checked were accurate, and the book is agree- 
ably free of typographical mistakes. I am less enthu- 
siastic about the indices, which are divided into “prey 
species index” and “subject index”, and the former is 
further subdivided within the index itself. ’'ve never 
been able to comprehend the rationale for split indices, 
but these seem adequate, if irritating. 

This book is a major contribution to the literature 
on this subject. It is a “must” for anyone working in 
this or allied fields, an important reference text, and a 
comprehensive, stimulating and thought-provoking syn- 
thesis of a very important area of animal behaviour. 


CLIVE E. GOODWIN 


| Queen Street Suite 405, Cobourg, Ontario K9A 1M8 
Canada 


2005 


For Love of Insects 


By Thomas Eisner. 2003. The Belknap Press of Harvard Uni- 
versity Press. Cambridge, Massachusetts. 448 pages. Paper. 


I remember well being almost overcome with wafts 
of formic acid after breaking apart an old black spruce 
stump during a stroll in the forests of eastern New- 
foundland. At the time I figured that I was the victim 
of an ant species of the subfamily Formicinae, ants 
that Eisner describes as “spray gun(s) on legs.” Such 
ants, each containing a formic acid gland, can eject a 
fluid that may exceed 50 percent formic acid. Put hun- 
dreds of such ants together in one spruce stump and 
the chemical barrage can be overwhelming. 

A self-described “incorrigible entomophile,” Thomas 
Eisner, Schurman Professor of Chemical Ecology at 
Cornell University, has written a feast of a book. In 
astonishing detail and delight, he describes a host of 
insects and arthropods and the myriad ways in which 
different species have evolved sophisticated chemi- 
cally-based means of predation, protection, mating, and 
a host of other interactions among insects or between 
insects and plants. 

Eisner’s early passion for the natural history of 
insects combined with his interest in natural chemistry 
led to his pioneering advances in the chemical ecology 
of insects and arthropods. Chemical ecology came to 
be his life-long passion as he delved into the secret 
lives of insects mediated by a host of chemical signals. 
The beginnings of potential research projects were often 
formulated during his favourite walks in nature, partic- 
ularly in the southern and southwestern United States. 
These initial observations often led to further sophis- 
ticated chemical studies on the identity and mechanism 


Book REVIEWS 


601 


of the chemical basis of insect behaviour. 

The book is a feast of stories about insects and their 
wonderfully adaptive chemicals that allow them to live 
and survive what is obviously a precarious existence. 
Each story is adorned with fascinating photos (many 
from the adept hands of Eisner himself) of a world sel- 
dom seen or appreciated by most of us. 

These stories of insects and their chemical ecology 
are wonders and gems in themselves. Of equal or even 
greater delight, is the sense of wonder and love com- 
municated by Eisner throughout the book’s pages. By 
self admission, the study of nature for this naturalist 
explorer can never disappoint him. His passion is infec- 
tious. His stories are peppered with the joy and exhil- 
aration of research and discovery. The reader is intro- 
duced not simply to facts and explanations, but to a 
host of graduate students, post-docs, and collaborators, 
including his wife Maria, who shared the joy. and 
delight of Eisner in his “love of insects.” 

Eisner ends his work by noting that without curiosi- 
ty, without passion for discovery, nature cannot endure. 
He laments our steady encroachment upon nature, 
upon the loss of boundaries. He notes, however, that 
ultimately, curiosity will be sustained — and that we 
will coexist with the living world. Can love of insects 
make a difference Eisner wonders? He ends this treas- 
ure of a book by admitting that he would like to believe 
that it does. I could not agree more. 


JOHN MCCARTHY 


P.O. Box 1238, (Station Main) Guelph, Ontario NIH 6N6 
Canada 


Rare Bird: Pursuing the Mystery of the Marbled Murrelet 


By Maria Mudd-Ruth, 2005. Holtzbrinck Publishers, 175 
Fifth Avenue, New York, New York 10010. 298 pages. 
US$16.29. 


This well-written environmental history book cap- 
tures a fascinating story: the Marbled Murrelet, a 
seabird nesting mostly in trees: one of the greatest 
environmental failures in the conservation history of 
globalization, western civilization and North American 
governmental stewardship. The book states that since 
1830 due to logging of the old-growth forest unsus- 
tainable activities in the Pacific Northwest has spared 
only 10% of the historic nesting habitat of the Marbled 
Murrelet. Other well-documented threats for Marbled 
Murrelets in this book, are fisheries (gill net by-catch) 
and oil pollution. That this species is covered by the 
U.S. Endangered Species Act (ESA) is a controversial 
topic because, in areas around Seattle and north of it, 
it is among the most abundant birds almost year round. 

The book reads well and can be recommended to 
naturalists, arm-chair explorers as well as to conser- 
vationists, ornithologists, Marbled Murrelet activists, 


and researchers. Its great strength is its documenta- 
tion of the recent history of Marbled Murrelet research 
in the United States, in California, Oregon and Wash- 
ington. Also, an archeological and native view from 
the Haida and Tlingit tribes is presented, including 
fascinating stories about specimen and egg collections 
now distributed world-wide (e.g. in museums of Lon- 
don and Vienna). Accounts of famous explorers and 
early naturalists in “Marbled Murrelet land” such as 
Latham, Gmelin, Banks, Pennant, Lever, Cantwell, 
Dowell and Jewett are included as well. 

The text reads like a detective story: The toenail of 
the Marbled Murrelet is as well covered as the first 
official nest found in 1974 near San Francisco by H. 
Foster. Other fascinating details cover a Canadian nest 
encountered 1953 by W. Feger, how S. Sealy initiated 
the first Marbled Murrelet studies in North America, 
and Russian scientist S. Kishchinski’s theory of juve- 
niles leaving the nest to the remote ocean by simply 
following a stream. Insights are provided on how the 
International Council for Bird Preservation accepted 
Marbled Murrelets as a conservation issue of global 


602 


importance. And the reader will enjoy to learn why 
Ridgway’s colour code matters for Marbled Murrelets, 
why K. Nelson converted from studying woodpeckers 
to Marbeled Murrelets, about T. Hamer’s infamous 
eggshell surveys, and the pioneering work by S. Sealy’s 
student H. Carter. But it also deals with a heroic des- 
cription of the many unfunded Marbled Murrelet pio- 
neers that went to promote science in the public con- 
servation battle, or simply to advance ornithological 
knowledge. The book makes clear that one nesting tree 
alone can fetch $100 000, resulting in huge sums be- 
ing at stake when many hectares of forest are up for 
public discussion for potential logging. These high val- 
ues explain the subsidized heli-logging of Marbled 
Murrelet habitat on steep slopes at high elevations. 

The book handles the conservation failures for the 
Marbled Murrelet well, specifically the ones from the 
many governmental agencies involved (the reason why 
this book had to be written in the first place). Author 
Mudd-Ruth presents these shortcomings with great 
skill, such as the early failures of the U.S. Fish and 
Wildlife Service (did not carry out surveys) and U.S. 
Forest Service (ignored habitat inventories). But besides 
the notorious Bureau of Land Management and even 
the National Park Service, one could easily extend 
these shortcomings to the suite of state agencies such 
as Departments of Game, Natural Resources, Parks and 
Recreation, and certainly Forestry. And since oceans 
are directly involved in this topic, the National Insti- 
tute of Marine Fisheries, the National Oceanic and 
Atmospheric Administration, the National Sanctuaries 
and the U.S. Navy all get to carry the blame as well. 
The Canadian side involves at least the Canadian Wild- 
life Service, the Department of Fisheries and Oceans, 
the British Columbia Ministries of Forestry and Envi- 
ronment, Parks Canada, and the Mineral Management 
Service. The Marbled Murrelet swims with steadily 
declining numbers in all of these “governmental wat- 
ers”, as well as in the industrial and private ones. And 
almost all of these agencies have a noted Marbled 
Murrelet conservation history, and some, to this very 
day, are doing nothing of relevance to resolve the Mar- 
bled Murrelet problem. 

It becomes clear from this book that these institu- 
tions do not provide relevant public guidance and high 
quality data for sustainable decision-making, and ad- 
dressing the inherent conflict between endangered 
species, economic growth and development. Are the 
existing administrative structures and the reward and 
promotion systems for the staff of these agencies really 
able to manage such problems at all? 

The author mentions a lot the Pacific Seabird Group 
(PSG), which is involved as a lobby and expert group 
for Marbled Murrelets. However, the PSG is not a legal 
governmental body and can only compile and provide 
suggestions. To this very day we have no high quality 
old-growth inventory layer relevant to Marbled Mur- 
relet habitat and sustainable management of the pre- 
cious forest; nor do we even have any relevant studies 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


and data collected throughout its range on the food 
items of Marbled Murrelets; e.g., Sand Lance. As this 
powerful book outlines, the Pacific Northwest Forest 
plan created 1994 by the Clinton administration has 
improved the Marbled Murrelet management at least 
for the states of the lower 48. 

The book makes the tensions around the Marbled 
Murrelet situation explicitly clear: In 1989 the Mar- 
bled Murrelet was suggested as a candidate species 
for the ESA. However, it took three more years. When 
the Audubon Society of Portland sued the U.S. Fish 
and Wildlife Service the bird eventually received a 
status in 1992 federally listed as a threatened species; 
when it was the recovery process could start. The status 
review of this endangered species is due every five 
years, but as the reader will learn, it has been delayed 
from 1997 til 2002 when eventually a timber industry 
group took the government to court. Another classic 
‘“MaMu” legal case is presented with the influential 
court case from 1993 “Marbled Murrelet vs. Pacific 
Northwest” where EPIC (Environmental Protection 
Information Center) sued Pacific Lumber. 

Further, as well-presented in this book, Marbled 
Murrelets live in a complex habitat; e.g., where one of 
the biggest marihuana grower industries in the western 
world makes its (illegal) living. Raven populations in 
the Pacific Northwest have increased 15-fold: a human 
subsidized Marbled Murrelet nest predator. Habitat 
fragmentation is severe, and the outlook is very bleak, 
as the loss of Redwoods shows for instance. 

This book could have been improved if the Russian 
and Canadian populations were explained and elabo- 
rated on in more detail: their story is as exciting and 
as full of tensions as the U.S. one and inclusion would 
provide a complete context. Most relevant Canadian 
researchers and very influential players are not named 
and elaborated on in this book, such as I. Manly, V. 
Bahn, G. Kaiser, A. Derocher, F. Cooke, A. Burger. 
This book also has no maps (but nice black and white 
figures by Paul Jones). It was the Canadians who solved 
one of the biggest technical problem in the study of 
this bird: being able to catch it. The brood patch story 
presented in this book was already revised when this 
book was written and by Canadian researchers who the 
author quotes. The Canadian Clayoquot Sound (just 
north of Seattle), the location of a major environmental 
conflict and Canada’s largest mass-arrest in history, 
is hardly mentioned at all. 

The book explains that more than 3000 trees have 
been climbed in order to find the elusive nests, but less 
than 61 known nests are reported in this book (likely, 
one has climbed trees and searched in the wrong field 
sites and habitats). This leads me to a more serious flaw 
in this book. The author chose to ignore the known 
nests and findings; e.g., as presented at PSG meetings 
at which the author personally was present and referred 
to and cites in the text. Marbled Murrelet nests are 
actually located across elevations and on steep slopes, 
which will modify the conservation pressure on habi- 


2005 


tats. The reader is advised to adjust the nest site image 
presented in this book at the following website by the 
Center for Wildlife Ecology http://www.sfu.ca/biology/ 
wildberg/mamuweb/welcome.htm where over 120 nests 
are presented to the public. 

Unfortunately, this book does not document that 
“low elevations” alone does not sufficiently describe 
the nesting habitat. It does not deal with how the myth 
of low elevation old-growth forest evolved in the first 
place, and why this myth remains active within the 
conservation community to this very day. The book 
reports an uncommented figure for 10s of millions of 
Marbled Murrelets in Alaska alone; this must be way 
too high by the factor 10. 


BOTANY 


Canada’s Forests: A History 

By Ken Drushka. 2003. Forest History Society Issues Series. 
McGill-Queen’s University Press, Montreal and Kingston. 
97 pages. Paperback. $12.95 Can. 


This treatise was written by a journalist, now de- 
ceased, who had considerable practical experience in 
the forest industry, having worked as a logger in 
British Columbia. It is highly readable, being target- 
ted at the nonspecialist, with few references or sup- 
porting data. Even so, it provides considerable food for 
thought in a compact format. The treatise presents a 
generally positive picture of Canadian forests and the 
forest industry. Ken Drushka states his premises boldly: 
“Canadian forests still exist virtually intact. Since Euro- 
pean colonization ... only about 6 percent of Canada’s 
forest land has been converted to other uses” (page vii). 
Moreover, “it is inaccurate and misleading to claim 
that the country’s forests are in the process of being 
destroyed, devastated, decimated, or damaged beyond 
recovery” (page 82). On the contrary, Drushka con- 
siders that, by and large, “they are whole and healthy” 
(page 82). This is certainly a view markedly at variance 
with that usually presented through the popular media. 

Drushka develops his views in five chapters, focus- 
sing on “The Canadian Forest”, “Early Forest Use”, 
“Industrialization of the Forests”, “The Rise of Forest 
Conservation”, and “Sustainable Forest Management”, 
which includes a brief survey of the state of Canada’s 
forests as of year 2000. The survey covers a lot of 
ground (pun intended!) and, as might be expected, 
the treatment of topics is somewhat uneven and often 
cursory. I found the summary of the postglacial 
assembly of “The Canadian Forest” especially unsat- 
isfactory. Also in this opening chapter, Drushka sets 
the scene by identifying and describing eleven mod- 
ern forest regions across Canada, although only ten 
are shown on the accompanying map. Following this 
introduction, his historical survey in the next four 
chapters 1s basically chronological and sequential. 

Drushka notes that the Canadian landscape has been 
home to people throughout postglacial time and that 


BooK REVIEWS 


603 


No doubt, the peculiar efforts by the current U.S. 
government administration trying to de-list this species 
from the ESA and against all efforts of most biologists 
will keep this well written and very readable book in 
high demand. It’s the first of its kind for the Marbled 
Murrelet, and a great description for the state-of-the- 
art of the environment and conservation in arguably 
one of the most powerful countries in the world. 


FALK HUETTMANN 


Biology and Wildlife Department, Institute of Arctic Biology, 
419 Irving I, University of Alaska, Fairbanks, Alaska 
99775-7000 USA 


Aboriginal people “had an often-significant impact 
on the forests” (page 17). He points out that Aboriginal 
people used fire as a land management tool. I found 
it refreshing to see an environmental history which 
acknowledges that Canada had a human history before 
European settlement. Drushka identifies the first major 
impetus for sustained forest use as the establishment 
of the fishing industry in Newfoundland in the 1500s. 
Interestingly, he does not highlight the fur trade as a 
major reason for forest clearance, pointing out that 
permanent settlement by Europeans was actively dis- 
couraged by the trading companies. He sees the main 
phase of European settlement, beginning in the sev- 
enteenth century, being associated with “serious and 
mostly negative impacts on the forests” (page 23), as 
land was cleared for agriculture and wood was used 
as the principal fuel for winter heating. Commercial 
exploitation of the forests began in the late seventeenth 
century, propelled by warfare and conflicts that stim- 
ulated Britain particularly to turn to Canada as a source 
of supply for timber. The exploitation front moved 
westward and inland from the east coast to interior 
Canada, primarily along the major waterways, follow- 
ing a similar pattern to settlement. Drushka traces the 
establishment of administrative policies that made the 
forestry industry a major source of government rev- 
enue, especially later at the provincial level. 

In the third chapter, Drushka examines more close- 
ly the industrialization of the forest industry through 
the latter part of the nineteenth century. He considers 
that the spread of transportation networks, especially 
the railway, had a triple effect on the forest industry. 
First, railway construction consumed large amounts 
of timber. Second, better transport networks encour- 
aged the spread of lumbering to previously economi- 
cally inaccessible regions. Third, the wood-burning 
steam locomotives themselves demanded large quan- 
tities of timber as fuel. Clear-cutting became common. 
Drushka points out that the spread of lumbering went 
hand-in-hand with devastating wildfires that spread 
through the remaining slash on clear-cuts, often start- 


604 


ed by sparks from various industrial machinery. Drush- 
ka notes a prevailing lack of concern with the reduc- 
tion in area of forest: “Well into the twentieth century 
the sentiment was still widespread that forests were an 
impediment to development and settlement, and their 
eradication was acceptable, if not desirable” (page 37). 
Nevertheless, by turn of the twentieth century, the re- 
duction in forest land had become so marked that it 
was starting to raise alarm in some quarters, especially 
in government circles. This awareness marks the begin- 
ning of the rise of the forest conservation movement. 

This movement sprang from “a desire to maintain 
forests in perpetuity” (page 61) and was part of a broad- 
er conservationist movement, driven by social and ide- 
ological trends. Drushka traces the practical application 
of forest conservation to the establishment of forestry 
schools at universities and founding of various forest 
societies. The overall result was the “professionalisa- 
tion” of forestry and the development of a cadre of 
scientifically-trained foresters. Canadian forest con- 
servation policies therefore were rooted in a belief 
that management had to be science-based and, as far 
as possible, separate from the political process. Drush- 
ka points out that in Canada forest conservation was 
essentially utilitarian and not preservationist, under- 
pinned by the belief that the forests were there to be 
used, albeit within limits. Hence, throughout various 
legislative and regulatory initiatives has run the idea 
of “sustainable yield”. However, Drushka argues that 
all these policy approaches since the Second World 
War have focussed on sustainable yield of timber only, 
while other valuable and sustainable aspects of forest 
lands, such as water quality and biodiversity, have not 
been included. The conclusion is that an overall com- 
prehensive approach was lacking, which set the stage 
for the conflicts of the last few decades. 

In the last chapter, Drushka examines forest man- 
agement in the closing decades of the twentieth cen- 
tury. This was a time of considerable conflict and re- 
thinking. It had become apparent that previous estimates 
of “sustainable yield” were not in fact sustainable, 
partly because replanting was not keeping up with 
extraction, partly because of variations in growth, and 
partly because forest inventories were not in place to 
allow for a realistic assessment of what was “sustain- 
able’. Yet the forest industry was trying to respond to 
increasing demand for forest-based products, such as 


ENVIRONMENT 


The Natural History of Bermuda 


By Martin L. H. Thomas, First Edition, 2004. Bermuda 
Zoological Society, P.O. Box FL145, Flatts, FL BX, 
Bermuda. 


Islands are the Rosetta stones of evolution. They also 
contain some of the most vulnerable natural habitats 
on the planet and many species at risk. These two fac- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


pulp and paper. At the same time, there were increas- 
ing demands being placed on forests from other users, 
including recreationalists and environmentalists, who 
placed different values on the land. As Drushka notes, 
clashes between various groups of users were becom- 
ing more common. In this context, I think many read- 
ers from western Canada will remember the blockades 
of logging roads and protesters being removed by 
police. Although Drushka mentions competing uses, 
he does not examine any in detail, and this superficial 
treatment might leave some readers wondering why 
these conflicts were so bitter. Drushka does indicate, 
however, that these clashes were highlighting incom- 
patibilities between uses and leading to greater polar- 
isation. Increasingly powerful, vocal, and articulate 
interest groups were lobbying for a preservationist 
approach to forest lands, partly through advocacy for 
the establishment of parks and other types of protected 
areas where the forest industry would be excluded. 
The result, concludes Drushka, was that forest man- 
agement and planning could no longer be the purview 
just of provincial forest services. The process had to 
be opened up. Drushka identifies a new approach to 
management, especially through the 1990s, called 
“holistic” forestry or “sustainable forest management’, 
which involves “a broadening of the concept of sus- 
tained yield to include all components of a forest” 
(page 69). He sees this change in philosophy as an 
impetus for the development of the Canada Forest 
Accord (1998), which acknowledges that forest eco- 
systems should be managed to benefit a broad spec- 
trum of users. One of the more interesting outcomes 
of this shift in perspective has been the development 
of various certification programs. Drushka observes 
that such programs provide an incentive for forest 
companies to practice good management. Neverthe- 
less, as Drushka’s survey shows, sustainable forest 
management is clearly more demanding than the old 
style “cut and move on” approach, requiring greater 
flexibility and adaptability on the part of the forest 
industry and a recognition that, no matter how vast 
they may seem, the forest lands are a finite resource. 


ALWYNNE B. BEAUDOIN 


Royal Alberta Museum, 12845-102"¢ Avenue, Edmonton, Al- 
berta TSN OM6 Canada 


tors make them of great interest to naturalists and ecol- 
ogists. These themes are explored in intimate detail in 
The Natural History of Bermuda. 

Bermuda is an archipelago of over 100 islands in the 
mid-Atlantic due south of Nova Scotia and 965 kilo- 
meters east of Cape Hatteras. Having a surface area 


2005 


of only 55 square kilometers, no place in Bermuda is 
more than a kilometer from the ocean. It has a sub 
tropical climate due to its situation in the Gulf stream, 
and a fascinating geological history. Some 100 000 
years old, Bermuda is a seamount with a limestone cap 
and red soil layers originating from sand blown from 
as far away as the Gobi Desert. It has a classic karst 
landscape perforated by caves and sink holes. 

With 63 000 inhabitants it counts as one of the most 
densely populated places on earth, much exacerbated 
by an annual influx of 600 000 tourists attracted by 
its picture post card beauty. Humans have been coming 
to Bermuda since the 1500s when it was first sighted 
by Europeans, with permanent settlement dating from 
the 1600s. The island’s terrestrial habitats were quick- 
ly transformed; originally covered by forest these were 
sadly depleted by ship building and other industries 
and today one finds only tiny remnants. The original 
inhabitants included a flightless rail and a “crow” rap- 
idly exterminated by the new arrivals. In the author’s 
words, “we may never know what delicate animals 
and plants were eradicated.” 

The author repeatedly returns to this theme, describ- 
ing how humans have, intentionally or not, altered the 
Island’s ecology. Boars, introduced by sailors prior to 
permanent settlkement undoubtedly finished off the 
flightless rail and other vulnerable fauna. Two other 
examples tell the tale: Yellow-crowned Night Herons 
were successfully reintroduced to control an accident- 
ly introduced land crab which had the bad manners 
to dig golf ball sized holes in golf courses, making a 
“hole in one a certainty”. Less successfully, anoles 
were introduced to control an accidently introduced, 
destructive fruit fly but ended up eating ladybugs which 
had in turn been introduced to control an accidently 
introduced scale decimating native trees. Great Kiska- 
dees, introduced to control the anoles, ended up eat- 
ing the endangered Bermuda skinks. And so it goes. 

Like all remote oceanic islands, Bermuda has a lim- 
ited terrestrial biodiversity, consisting mainly of species 
that can be transported over long distances by the wind 
or ocean currents. This paucity is illustrated by the con- 
trast between the number of native vascular plants, 156, 
and the number of marine fish, 423. Native inhabitants 
include birds, insects and plants originating from North 
America and the Caribbean: the marine life is distinct- 
ly Caribbean, sharing many of the same colourful 
reef species. Some of the arrivals have diverged suffi- 
ciently from their ancestors to become endemic species 


Saskatchewan Uncommon Views 


By John Conway. 2005. The University of Alberta Press, 
Edmonton. 135 pages. Illustrated. $29.95. 


The results of John Conway’s photographic forays 
across Saskatchewan are certainly “uncommon”. 
Sometimes strikingly beautiful, always minimalist, his 
photographs elicit reflection, nostalgia, even humour. 


BOOK REVIEWS 


605 


or subspecies, for example the Bermuda Skink is 
thought to have evolved from a shared ancestor with 
the North American Five-lined Skink. Perhaps the most 
celebrated member of the Bermudian assemblage is 
the Cahow, or Bermuda Petrel; originally known only 
from fossils it was famously rediscovered in 1951 
when a scant few pairs were found breeding. 

All of this is set out by the author in 21 twenty chap- 
ters variously focused on key environments (reefs are 
the largest ecosystem, fresh water ecosystems the 
smallest) or groups of animals. The book is exten- 
sively illustrated with colour photographs but avoids 
being a “coffee table” book by the detailed narrative 
which introduces the reader to each theme and des- 
cribes the key inhabitants and processes involved. At- 
tractive maps are found on the front and end pieces. 
Occasionally the text reads like a catalog, but there is 
enough analysis and sufficient interesting observations 
that the reader’s interest should be rekindled. It also 
reads from time to time like a university lecture, which 
is not surprising given that the author is a university 
lecturer. Because each chapter is rather self standing 
there is a fair amount of irritating repetition which 
could have been reduced through a final editing ses- 
sion. A few proofing errors are present such as men- 
tion of the “Blade-headed Gull’, surely a cut above the 
usual lariid, and the geographically confused Antiguan 
Anole which apparently hails from Barbados as op- 
posed to the Barbadian Anole transported from Antigua. 
Whiie most groups are well covered, marine mollusks 
and sea shells receive scant attention which is curious 
given the level of interest these attract. 

The final chapter looks to the future. Exports of the 
valuable endemic Bermuda Cedar were banned as early 
as 1657, however, systematic conservation measures 
only emerged much more recently. Marine ecosys- 
tems are relatively well protected, however, only 7% 
of the land surface receives official protection. A pro- 
tected species act was introduced in 2003 and there 
are prohibitions on the importation of new species. 
Environmental education, particularly of youth, is a 
key priority and it is to be hoped that this handsome 
book will help especially as it is accompanied by a 
CD, making it an “electronic book.” If you have an 
interest in island biogeography, or just want to dream 
about your next escape from a Canadian winter, this 
book will be of interest. Magra est 
Permanent Mission of Canada, 5 avenue de |’ Ariana, Gene- 

va, Switzerland 


While this is not a book on natural history, it will ap- 
peal to many readers, especially to serious practition- 
ers of photography and to residents of Saskatchewan 
steeped in its history and geography. Superficially, 
the photographs confirm the popular belief that Sas- 
katchewan is flat, dull and colorless. But a closer 


606 


scrutiny (St Denis hills, for example page 43, or cat- 
tle and hay bales in a snowy landscape, page 5) sug- 
gests the beauty he could have depicted had that been 
his primary intention. In addition to the photos, the 
book contains three thoughtful essays by well known 
Saskatchewan writers: Sharon Butala, Helen Marzolf 
and David Carpenter. A careful reading of these essays 
is likely to result in a deeper appreciation and under- 
standing of Conway’s art and sensibility. 

Essentially, Conway shows what the prairie once was 
and what has since happened to it and to the dreams 
of thousands of farm families who immigrated to the 
West in the first decades of the twentieth century. His 
photos suggest a variety of reactions to a land that can 
be both inviting and cruel. Small towns are disappear- 
ing; many farm homes sit forlorn and abandoned; the 
larger urban centres reflect a new way of life, essen- 


MISCELLANEOUS 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


tially industrial and commercial, isolated from the land 
which nurtured aboriginals and early settlers. How- 
ever, several of his pictures reveal the tenacity and 
spirit of the people still living in rural Saskatchewan: 
the remains of a bicycle mounted on a fencepost in 
imitation of modern art (page 13); a roadside sign: 
Shirl’s Upholstery, standing in heavy grass. Conway’s 
caption: “Shirl has done upholstery out of a mobile 
home for twenty five years.” 

A handsome book, reasonably priced, Saskatche- 
wan Uncommon Views is a significant achievement, 
particularly fitting since it appeared in the province’s 
centennial year. 


J. FRANK Roy 


912-606 Victoria Avenue, Saskatoon, Saskatchewan S7J 0Z1 
Canada 


Manly Hardy (1832-1910): The Life and Writing of a Maine Fur-buyer, Hunter, and Naturalist 


By William B. Krohn. 2005. Maine Folklife Center, 5773 
South Stevens Hall, University of Maine, Orono, Maine, 
USA 04469. 343 pages. $24.95 hardcover, $19.95 paper. 


Manly Hardy documented more about Maine’s wild- 
life during the last half of the 19" century than any 
other writer. He was “a keen and reliable observer ... 
a faithful recorder,” and a “widely capable, fiercely 
independent, highly intelligent and positive man.” 
Krohn demonstrates “just how dynamic Maine’s nat- 
ural environments were during much of the nineteenth 
century.” This book was possible because the author 
had access to the documents collected by two unrelat- 
ed people: Fannie Hardy Eckstorm, Hardy’s first child, 
and the late Dr. Ralph S. Palmer. In addition, Palmer 
was responsible for much of the annotated bibliogra- 
phy which appears at the end of the book. 

The first 78 pages, written by W. B. Krohn, deal 
with Manly Hardy’s life. Hardy was a complex per- 
son with many seemingly contradictory qualities. A 
small and sickly child in the small town of Brewer, 
Maine, he became religious as an adolescent, and stud- 
ied Greek and Latin in a private school taught by a 
minister. In his twenties he became a hunter and trap- 
per who regularly took life for food and fur, yet he 
loved pets and could be sentimental about animals. 
When Hardy was 32, he inherited from his father the 
family fur business; he handled over half a million 
dollars worth of furs during his career, the largest fur 
business east of the Rocky Mountains. 

When Hardy was growing up, Indians outnumbered 
white people in Brewer. Hardy’s approach was unusu- 
ally enlightened for his time; hunting and traveling 
with them, he learned their ways. He respected their 
knowledge and skills. They confided in him and he 
guarded their secrets. 


Krohn deals mainly with Hardy’s interests outside 
of his business. Hardy amassed a collection of over 
3000 birds; he traded Passenger Pigeons from Maine 
to obtain some of the specimens. He wrote at least 
150 articles about nature, over half of which appeared 
in Forest and Stream. Ernest Thompson Seton (some 
of whose delightful sketches are reproduced in this 
book) considered Hardy’s writings to be among the 
most useful information in print; he quoted Hardy 72 
times in his monumental, four-volume Lives of Game 
Animals (1925-1928). Ralph S. Palmer cited Hardy 
21 times in Maine Birds (1949). 

The main portion of the book, 203 pages, repro- 
duces some of Hardy’s most interesting articles, in- 
cluding two accounts of long winter trips in the Maine 
woods. The mammal articles selected by Krohn con- 
cern four species that are now extinct or nearly so in 
Maine, the woodland caribou, wolf, cougar and sea 
mink, plus others of special interest, the lynx, bobcat, 
fisher, moose, beaver, river otter, and porpoise. Hardy’s 
well-written accounts are still of interest today, offer- 
ing the best available comparison with what things 
were like more than a century ago. The final 32 pages 
give an annotated bibliography of Hardy’s other bird 
and mammal vignettes. I found only one minor error, 
made by Hardy, who misspelled the surname of Rod- 
erick Ross MacFarlane [not MacFarland]. 

For those with an interest in the history of fur-bear- 
ing mammals in the days when Maine’s woods were 
in a pristine state, this book is a treasure. It will be of 
nearly equal interest to residents of adjacent New 
Brunswick. 

C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N O0J8 
Canada 


2005 


BooK REVIEWS 


607 


Audubon in Edinburgh and his Scottish Associates 


By John Chalmers. National Museums of Scotland, Edinburgh. 
2002. 157 pages. $34.95 U.S. $70.00 Can. Available from 
Codasat Canada Ltd., #1, 4335 West 10" Ave., Vancouver, 
British Columbia V6R 2H6 Canada. 


John Chalmers, an orthopedic surgeon in Edinburgh, 
has filled a void with his careful documentation of 
archival letters and files dating back more than a cen- 
tury and a half. Although most members of the general 
public know the name of Audubon, few historically- 
minded ornithologists realize that Edinburgh, Scotland, 
was Audubon’s favourite among all the world’s cities. 
Audubon had received little encouragement in Amer- 
ica, or in London or Paris, but Edinburgh grew to love 
him during the 22 months he spent there during his 
six visits, two of which lasted for six months each. 
Audubon’s fame and his success date from these six 
visits. Chalmers includes twelve pages of biographi- 
cal profiles of men who interacted with Audubon. 

Audubon, the rough but charming and talented back- 
woodsman, was unexpectedly well received by emi- 
nent people in what was then the world’s leading city 
for science endeavours. Members of the intelligentsia 
wined and dined him. The august Wernerian Society 
welcomed him. Thirteen of Audubon’s early scientif- 
ic papers were published by four different Edinburgh 
scientific journals. William McGillivray, in particular, 
went out of his way to help him with the scientific 
aspects of ornithology. The kindness of such savants 
never ceased to amaze Audubon. 

Audubon’s Birds of America was extremely expen- 
sive to produce. The first 10 plates were engraved by 
W. H. Lizars in Edinburgh, though publication later 
switched to London. In total, there were 87 parts, which 
sold at two guineas each, for a total cost of £187, con- 
taining 435 plates and text. Chalmers provides a list 
of the present whereabouts of the 23 sets that were 
sold in Scotland, as well as a list of exactly fifty places, 


A History of Devonshire Ornithology 


By David G. Jenks. Isabelline Books, 2 Highbury House, 8 
Woodlane Crescent, Falmouth TR11 4QS, Cornwall; e- 
mail: mikann@beakbook.demon.co.uk. FAX 0870 051- 
6387. 2004. Hardcover. 477 pages. £53 plus postage. 


Devon is one of the most beautiful and interesting 
of the many British counties. David Jenks planned to 
write its ornithological history as a chapter for Birds 
of Devon, but when that project fell through he com- 
pleted an entire book on the history alone. And what 
a book! Full of detail, printed on high quality paper, it 
is a sumptuous book, but almost too heavy to hold for 
someone who reads a chapter each night at bedtime. 

As the Foreword explains (page xi), thoroughly- 
researched historical accounts such as this help us 
“understand the origins and raison d’étre of the mod- 
ern nature conservation movement.” The first chapter 


including 43 private residences, frequented by Audu- 
bon; all but 11 of these buildings are still standing. 

Ironically, not one of the 14 complete sets of Birds 
of America sold in Edinburgh has remained there; the 
cash-strapped University of Edinburgh sold its set at 
auction in New York in 1992 for $4.1 million. There 
is an extant copy in Paisley, coincidentally the home- 
town from which Alexander Wilson had left in dis- 
grace for America in 1794. 

This book combines an informative text, helpful 
footnotes, and magnificent illustrations, including re- 
productions of some of Audubon’s finest bird paint- 
ings. Readers will marvel, as Audubon did, at the extent 
to which Edinburgh launched his career. Among in- 
numerable pleasures are the spontaneity and charm 
of Audubon’s letters to his wife, back in America. 

I detected few errors. Although Chalmers is correct 
in saying that Alexander Wilson, a transplanted Scot 
in America, named the Eskimo Curlew in honour of 
Audubon in 1813, he fails to appreciate that J. R. Forster 
in 1772 had pre-empted Wilson by naming the species 
from a specimen collected by Humphrey Marten at 
Albany, on Hudson Bay, in 1771. Elliot Coues is mis- 
spelled Cowes. The index is sadly incomplete, hardly 
excusable now that indexes can be compiled unerr- 
ingly and almost automatically by highlighting each 
name in the text. 

This sumptuous, 228-page, beautifully-illustrated, 
sturdily-bound paperback is necessarily expensive. It 
would be an elegant gift for anyone interested in Au- 
dubon, the primacy of Edinburgh during Scottish en- 
lightenment, or the history of ornithology. One hopes 
it can be:purchased by every major library in North 
America and Europe. 

C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N OJ8 
Canada 


deals with the prehistoric record — skeletal remains of 
birds found in seven different caves or deposits. These 
caves are mapped, but readers are handicapped by the 
lack of a map of Devon showing the many other place 
names mentioned. 

North Americans can only envy the additional cen- 
turies of scattered historical information available in 
England. The Isle of Lundy, off the north shore of 
Devon, features prominently throughout. The first 
Lundy entry concerns the gannet colony and nesting 
peregrines found there in 1274 AD. 

There is a sordid side to most of the accounts before 
1900. So-called scientific interests were served by 
shooting a bird, then almost the only way to identify it. 
Physicians and country vicars alone seemed to have 
“the leisure, the academic training and the opportunity 


608 


to become well-informed naturalists” (page 135). More 
often than not it was the local minister of the Church 
of England who was guilty of massive slaughter of 
almost anything that flew. Jenks tells of the zealous 
collector who, while walking without a gun, saw an un- 
familiar bird, knocked on the door of the nearest cot- 
tage, borrowed a gun from the stranger, and shot what 
proved to be a beautiful Night Heron in full breeding 
plumage. 

Then there was the mercenary side of bird-killing. 
Village markets throughout Britain were replete with 
waders, marsh birds, even song birds, all of them to be 
purchased for the table. They fetched a surprisingly 
high price. 

Quite apart from culinary sales, both birdskins and 
eggs were also available. Those interested in the dis- 
tribution of bird species were apt to go to their local 
market and purchase the rare birds available for sale. 
In 1878 a Gyrfalcon skin sold for 50 shillings and a 
Golden Eagle egg for 25 shillings; a Passenger Pigeon 
skin cost only 6 shillings. As evidence of the extent 
of this trade, the relatively small town of Barnstaple 
supported two full-time bird-stuffers. 

We can learn from some of the mistakes made in 
Devon. Many specimen records were made worthless 
by inadequate preservation, missing date and locality 
labels, and some were unknowingly brought by ship 
from offshore. 

Throughout the book there is evidence of friction 
between those who collected birds’ eggs and study 
skins and those who used the new field glasses to ob- 
serve birds in the wild. The latter slowly and gradually 
came into the ascendancy after 1900. 

We meet many interesting men, but I will mention 
only two. Colonel George Montagu, who wrote his 
landmark Ornithological Dictionary in 1802, was 
Devon’s first important ornithologist; he sorted out the 
differences between the Montagu Harrier and the Hen 
Harrier, thus having his surname applied to the former. 
The first imperfect beginnings of a book to be devoted 
entirely to Devonshire birds was written by Andrew 
Tucker in 1809; only two of the proposed 24 parts ap- 
peared in print. Surprisingly, it was not until 1916 that 
Catharine Hodgins became the first Devon woman to 
contribute bird observations for publication. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Lists of spring migration dates compiled from 1840 
to 1845 indicate that birds now arrive much earlier, 
evidence of climatic change. Some notable sightings 
include a flock of eight Great Bustards in 1870, never 
to be repeated. The Eurasian Collared-Dove first ap- 
peared in Devon, on Lundy in May 1961, only six years 
after the first pair had appeared in Norfolk; thus the 
spread across Europe preceded its recent spread across 
North America. 

Equally interesting are occasional accounts of un- 
usual happenings. In the winter of 1928-29, Water Rails 
in desperation entered into houses to take bread off 
tables. A peregrine found dead near its eyrie on Lundy 
in 1963 proved by retrospective analysis, once Derek 
Ratcliffe discovered the causative role of DDT and 
other organochlorine compounds, to be “the most 
chemically-contaminated individual ever discovered.” 
(page 334). 

I found the final chapter, on bird-ringing, of partic- 
ular interest. David Lack, then a schoolmaster at 
Partington in Devon, prior to his distinguished career 
as a professor at Oxford University, ringed 119 adults 
and 121 nestling Robins on one leg and applied two 
coloured celluloid rings to the other leg. Many robins 
remained from year to year, the oldest persisting through 
all four years of his study. Not one was recorded out- 
side his study area of twenty acres. His famous book, 
The Life of the Robin, told what he learned about in- 
dividual robin territories and behaviour. 

Ringing changed forever when the first shipment 
of 100 mist nets arrived from Japan in early 1956. A 
formal ringing station, using these nets, opened at Slap- 
ton in 1960. Jenks, a ringer himself, concludes by say- 
ing (page 390) that “ninety years of ringing in Devon 
has provided us with more knowledge of the county’s 
birds than all other ornithological pursuits put togeth- 
et. 

In summary, this book constitutes a fine and lasting 
contribution to the history of ornithology. Learning 
about past excesses can give us perspective and per- 
haps help us to prevent similar but different errors in 
future. Its widest readership will undoubtedly be in 
Great Britain. 

C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8, 
Canada 


Mapper of Mountains: M. P. Bridgland in the Canadian Rockies, 1902-1930 


By I. S. MacLaren, with Eric Higgs and Gabrielle Zezulka- 
Mailloux. University of Alberta Press, Edmonton, Alber- 
ta. 2005. 295 pages. $39.95 Can. Cloth. 


This book, well written, artistically presented, and 
superbly illustrated, explains the history of photo- 
graphic mapping in the Canadian Rockies. Compli- 
cated technical processes are clearly explained. As one 
would expect from Ian MacLaren, the text is enlivened 
by the poetry of Robert W. Service, Bliss Carman 


and, surprisingly, the famous Scots prose author, Sir 
Arthur Conan Doyle. Serious readers will appreciate 
the detailed information in the copious footnotes. 
The subject, Morrison Parsons Bridgland (1878- 
1948), sometimes called Morris, but more often known 
as M. P., was exceedingly modest. There are no known 
photographs of him alone. Born on a farm on the 
northern outskirts of Toronto, M. P. studied engineer- 
ing at the University of Toronto, and then was hired 


2005 


as an apprentice land surveyor. He was remarkably 
fit, with incredible endurance; he once climbed 7600 
feet between 2 a.m. and 11 p.m. On another occasion 
he spent five hours ascending the glacier on Mount 
Purity, but was able to glissade down in 12 minutes. 
He was the first to climb to the peak of 55 different 
mountains, and took photographs from each. 

Difficult and dangerous climbs were routine. Con- 
tending with bears and snowstorms, and living fru- 
gally, M. P. became one of the foremost phototopo- 
graphical surveyors anywhere in the world, taking more 
photographs of higher quality than anyone before him. 
He carried a 35-pound camera up steep mountainsides 
in its mahogany box, fitted into a leather case. The 
photographs were exposed on fragile large glass slides. 
An assistant carried the transit instrument (theodo- 
lite) and the tripod. Since photographs at the standard 
f 32 stop required a shade over the lens, a filter, and 
exposures of up to three seconds, absolute immobility 
had to be achieved in spite of insecure footing and 
howling winds. On each mountain top, they built a 
cairn, then took exquisite care in leveling the camera 
to define the horizon and obtain photographs in every 
direction, but especially to show cairns on adjacent 
peaks, so that angles between them could be measured 
precisely. A tent with absolute darkness became the 
darkroom to develop a few test exposures each time, 
but the other exposed plates were carried down the 
mountain side and shipped “‘at great expense” to Ottawa 
for development the following winter. Precise mathe- 
matically, there was also an obvious artistic dimension 
to Bridgland’s work. In his Calgary office each winter, 
the photographs were used to construct accurate top- 
ographical contour maps. Including expenses for field 
and office work, Bridgland’s surveys were done for 
half the price of those completed by others, in Jasper 
park for only $4.20 per square mile. 

One of Bridgland’s important achievements was as 
a founder and chief mountaineer of the Alpine Club 
of Canada, helping with the society’s first three sum- 
mer camps and writing reports for the new Canadian 
Alpine Journal. Another memorable achievement, in 
1917, was his 97-page pamphlet, co-authored by Robert 
Douglas, secretary of the Geographic Board of Canada, 
Description of and Guide to Jasper Park. This well- 
written pamphlet (the design template for MacLaren’s 
book) opened with Sir Arthur Conan Doyle’s poem, 
the Athabasca Trail. It contained photographs of moun- 
tains, mammals and flowers, but most notably, Bridg- 
land’s striking photograph of Mt. Edith Cavell, named 


BooK REVIEWS 


609 


for the Canadian nurse shot by the German Army in 
World War I. The pamphlet sold for fifty cents and 
was reprinted many times; extant copies today fetch 
two hundred times the initial purchase price. A pack 
of six Bridgland topographical survey maps of Jasper 
park was also for sale, 90 cents unmounted, or three 
dollars mounted. 

On occasion, Bridgland was treated badly by the 
government bureaucracy in distant Ottawa. His party 
often worked seven days a week, and sometimes eight- 
een hours a day, but a civil servant in Ottawa, perus- 
ing Bridgland’s diary, noted that he had once taken 
one weekday off, and docked him a day’s pay. At 52 
years of age, after thirty years of service, Bridgland 
learned that his position of Surveys Engineer, Grade 4, 
had been abolished by the Bennett government. On 
12 May 1931 he was informed that his firing would 
be effective on 20 May, and that he would receive only 
three months of retirement leave. He never fully recov- 
ered from this treatment, but the Alpine Club of Canada 
awarded him the prestigious Silver Rope of Leader- 
ship. 

M. P. retired to Toronto where he died of leukemia 
in January 1948. His wife, Mary, survived another quar- 
ter century. His older son, Charles, graduated in elec- 
trical engineering from the University of Toronto. His 
younger son became Brigadier General Edgar Bridg- 
land of Canadian Aviation Development fame. 

In 1996, Jeanine Rhemtulla, a graduate student at the 
University of Alberta, began a study to re-photograph 
the vistas taken by Bridgland, to learn what changes had 
taken place. This Rocky Mountain Repeat Photography 
Project attracted additional helpers, including the two 
junior authors of this book. Photographic plates long 
thought to be lost were unearthed by deep historical 
sleuthing — and are now available for everyone to see 
on the web, http://bridgland.sunsite.ualberta.ca/index. 
html. The climax of this book is a series of photographs 
comparing Bridgland’s 1915 photographs with those 
taken over 80 years later. They demonstrate how gla- 
ciers have receded and forests expanded over that long 
time frame. 

Not only is this biography at its finest, the book is a 
perfect present for those interested in geography, moun- 
tains, climate change, or photography. Read it in prepa- 
ration for a visit to Jasper (especially), or Banff. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


610 


NEw TITLES 
+ Available for review * Assigned 


ZOOLOGY 


The Kingdon Pocket Guide to African Mammals. By Jon- 
athan Kingdon. 2005. Princeton University Press: 41 William 
Street, Princeton, New Jersey 08540 USA. 272 pages. $24.95 
Paper. 


* The Gulf of Alaska: Biology and Oceanography. Edited 
by P. Mundy (Ed). Alaska Sea Grant College Program, Uni- 
versity of Alaska, Fairbanks, Alaska 99775 USA. 214 pages. 
US$25. 


Snakes of the Americas. By R. Tipton. 2005. Krieger Pub- 
lishing P.O. Box 9542, Melbourne, Florida 32902-9542 USA. 
428 pages. US$94.50. 


A New World of Animals — Early Modern Europeans on 
the Creatures of Iberian America. By Miguel de Astia and 
Roger French. 2005. Ashgate Publishing Company, P.O. 
Box 2225 Williston, Vermont 05495-2225 USA. 276 pages. 
US$89.95 or £50.00 Cloth. 


Arrivals and Rivals: A Birding Oddity — A Year of Com- 
petitive Twitching. By Adrian Riley. 2005. NHBS Environ- 
ment Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 5XN 
United Kingdom £7.99, US$15. 


An Atlas of the Geographic Distribution of the Arvicoline 
Rodents of the World (Rodentia, Muridae: Arvicolinae). 
By G. Shenbrot and B. Krasnov. 2005. Pensoft Publishers, 
Acad. G. Bonchev Street, BI. 6, 1113 Sofia, Bulgaria. 350 
pages. EURO 77.90 Cloth. 


Retracing the Aurochs: History, Morphology and Ecology 
of an Extinct Wild Ox. By Cis van Vuure (University of 
Wageningen, the Netherlands). September 2005. Pensoft Pub- 
lishers, Acad. G. Bonchev Street, Bl. 6, 1113 Sofia, Bulgaria. 
424 pages. EURO 67.90 Cloth. 


A Field Guide Amphibians and Reptiles of Bali. By L. 
McKay. 2006. Krieger Publishing, P.O. Box 9542, Melbourne, 
Florida 32902 USA. No details available. 


* Band-tailed Pigeon: Wilderness Bird at Risk. By Worth 
Mathewson. 2005. Timber Press, 133 SW 2" Avenue, Ste. 450, 
Portland, Oregon 97204 USA. 196 pages. US$19.95, £14.99 
US$39.50 Cloth. 


Bengaliidae du Monde (Insecta, Diptera). By Andy Lehrer. 
2005 [In French, keys and summary translated in English.] 
Pensoft Publishers, Acad. G. Bonchev Street, Bl. 6, 1113 Sofia, 
Bulgaria. 192 pages. EURO 58.80 Cloth. 


Birds Britannica. By Mark Cocker and Richard Mabey. 2005. 
Random House of Canada Limited, 2775 Matheson Boule- 
vard East, Mississauga, Ontario L4W 4P7 Canada. 484 pages. 
$95.00 Cloth. 


Bird Coloration. Edited by G. Hill and K. McGraw. 2005. 
Harvard University Press, 79 Garden Street, Cambridge, 
Massachusetts 02138 USA. 496 pages. US$95. 


National Geographic Complete Birds of North America. 
By Jonathan Alderfer. 2005 National Geographic Society, 
1145 17th Street N.W., Washington, D.C. 20036 USA. 640 
pages. US$35.00 


Portraits of the Bison — An illustrated Guide to Bison 
Society. By W. Olsen. 2005. University of Alberta Press, Ring 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


House 2, Edmonton, Alberta T6G 2E1 Canada. 120 pages. 
Can. $39.85 Paper. 


Fauna Britannica. By Stefan Buczacki. 2005. Orion Publish- 
ing Group, Orion House, 5 Upper Saint Martin’s Lane, London, 
WC2H 9EA United Kingdom. £18.99, approximately $35/29 
Paper. 


Biomes of Brazil. An Illustrated Natural History. By F. 
Por, V. Imperatriz-Fonseca, and F. Lencioni. 2005. Pensoft 
Publishers, Sofia-Moscow. 208 pages. Euro 39.50 Cloth. 


Caterpillars of Eastern North America: A Guide to Iden- 
tification and Natural History. By David L. Wagner. 2005. 
Princeton University Press, 41 William Street, Princeton, New 
Jersey 08540 USA. 496 pages. US$29.95 Paper, $60.00 
Cloth. 


100 Caterpillars — Portraits from the Tropical Forests of 
Costa Rica. By J. Miller, D. Janzen and W. Hallwachs. Har- 
vard University Press, 79 Garden Street, Cambridge, Mass- 
achusetts 02138 USA. 300 pages. US$39.95. 


Field Guide to the Bumblebees of Great Britain and Ire- 
land. By Mike Edwards and Martin Jenner 2005. Ocelli 
Limited, 19 Church Street, Willingdon Village, Eastbourne, 
East Sussex BN20 9HR. 108 pages. £9.99 plus £1.50 shipping. 


* Crows — Encounters with the Wise Guys. By Candace 
Savage. 2005. Greystone Books #201-2323 Quebec Street, 
Vancouver, British Columbia VST 4S7 Canada. 113 pages. 
$27 Cloth. 


Dodo: The Bird Behind the Legend. By Alan Grihault 
2005, Imprimerie & Papeterie Commerciale Ltd Distributed 
by NHBS, 2-3 Wills Road, Totnes, Devon TQ9 5XN United 
Kingdom. 171 pages. £16.99. 


The Amphibians and Reptiles of El Salvador. By G. 
Kohler, M. Vesely and E. Greenbaum. 2006. Krieger Publish- 
ing, P.O. Box 9542, Melbourne, Florida 32902 USA. 248 
pages. US$49.50 Cloth. 


Fire Ants. By W. Tschinkel. 2005. Harvard University Press, 
79 Garden Street, Cambridge, Massachusetts 02138 USA. 
720 pages. US$95. 


+ Fire and Avian Ecology in North America — Studies in 
Avian Biology Number 30. Edited by Victoria A. Saab and 
Hugh D. W. Powell. Cooper Ornithological Society, c/o West- 
ern Foundation of Vertebrate Zoology, 439 Calle San Pablo, 
Camarillo, California 93012-8506 USA. 193 pages. US$18 
Paper. 


Fish, Fur and Feathers. By the Fish and Wildlife Historical 
Society. 2005. Federation of Alberta Naturalists, 11759 Groat 
Road, Edmonton, Alberta TSM 3K6 Canada. 


Hawks from Every Angle: How to Identify Raptors in 
Flight. By Jerry Liguori. 2005. Princeton University Press, 
41 William Street, Princeton, New Jersey 08540 USA. 144 
pages. US$19.95 Paper, $55.00 Cloth. 


Amphibians and Reptiles of the Honduran Mosquitia. 
By J. McCranie, J. Wilson, L. David and J. Townsend. 2006. 
Krieger Publishing, P.O. Box 9542, Melbourne, Florida 32902 
USA. No details available. 


2005 


Hummingbirds of Costa Rica. By M. and P. Fogden. 2006. 
Firefly Books, 66 Leek Crescent, Richmond Hill, Ontario 
L4B 1H1 Canada. 154 pages. $49.95. 


Identify Yourself: The 50 Most Common Birding Identi- 
fication Challenges. By Bill Thompson III and the editors 
of Bird Watcher’s Digest. 2005. Houghton Mifflin Company, 
215 Park Ave. South, New York, New York 10003 USA. 416 
pages. US$19.95 Paper. 


Insects: Their Natural History and Diversity. By S. Mar- 
shall. 2006. Firefly Books, 66 Leek Crescent, Richmond Hill, 
Ontario L4B 1H1 Canada. 704 pages. $95.00. 


The Other Insect Societies. By J. Costs. 2005. Harvard Uni- 
versity Press, 79 Garden Street, Cambridge, Massachusetts 
02138 USA. 602 pages. US$59.95. 


A Walk Around the Pond — Insects in and over Water. By 
G. Waldbauer. Harvard University Press, 79 Garden Street, 
Cambridge, Massachusetts 02138 USA. 258 pages. US$22.95. 


+ For the Love of Insects. By T. Eisner. 2005. Harvard Uni- 
versity Press, 79 Garden Street, Cambridge, Massachusetts 
02138 USA. 448 pages, US$16.95 Paper. 


Kaufman Field Guide to Insects of North America. By 
Kenn Kaufman and Eric Eaton. 2006. Houghton Mifflin Com- 
pany, 15 Park Avenue South, New York, New York 10003 
USA. 384 pages. US$18.95 Paper. 


In Search of the Ivory-Billed Woodpecker. By Jackson, 
Jerome. 2004. Houghton Mifflin Company 215 Park Avenue 
South, New York, New York 10003 USA. 294 pages. US$24.95. 


“The Grail Bird’’ — Hot on the trail of the Ivory-billed 
Woodpecker. Edited by Tim Gallagher, 2005. Cornell Univer- 
sity Press, Sage House, 512 East State Street, Ithaca, New York 
14850 USA. 288 pages. US$25.00 Cloth. 


Birds of Kuwait: A Portrait. By Abdullah Alfadhel. 2005 
Kuwait Environment Protection Society c/o NHBS 2-3 Wills 
Road, Totnes, Devon TQ9 5XN, United Kingdom. £29.95, 
approximately $56/45. 


Mako Sharks. By A. De Maddelena, A. Preti, and R. Smith. 
2005. Krieger Publishing, P.O. Box 9542, Melbourne, Florida 
32902 USA. 72 pages. No price available. 


Birding North Carolina. By Marshall Brooks, Mark Johns 
and Carolina Bird Club members. 2005. Globe Pequot Press, 
246 Goose Lane, P.O. Box 480, Guilford, Connecticutt 06437 
USA. 224 pages. U.S.$12.95 Paper. 


Marine Life of the Pacific Northwest. By A. Lamb and B. 
Handby. 2005. Harbour Publishing, P.O. Box 219, Madeira 
Park, British Columbia VON 2HO Canada. 400 pages. $69.95 
Cloth. 


North American Mushrooms: A Field Guide to Edible 
and Inedible Fungi. By O. Miller and H. Miller. 2006. The 
Globe Pequot Press, 246 Goose Lane, Box 480, Guilford, Con- 
necticutt 06437 USA. 592 pages. US$29.95, Can$35.25 Paper. 


Amphibians and Reptiles of Pakistan. By M. Khan. 2006. 
Krieger Publishing, P.O. Box 9542, Melbourne, Florida 32902 
USA. No details available. 


Birds of Washington. By T. Wahl, R. Wahl, W. Tweit, and S. 
Miodinow. 2005 Oregon State University Press, 102 Adams 
Hall, Corvallis, Oregon 97331 USA. 448 pages. US$65 Cloth. 


BOOK REVIEWS 


611 


North American Wildlife. By D. Jones. 2006. Firefly Books, 
66 Leek Crescent, Richmond Hill, Ontario L4B 1H1 Canada. 
304 pages. $29.95. 


The San Diego County Bird Atlas. By Philip Unitt. 2005. 
Ibis Publishing Company, 44970 Via Renaissance, Temecu- 
la, California 92590 USA. 645 pages. US$80.00 Cloth. 


+ Secret Weapons. By T. Eisner, M. Eisner, and M. Siegler. 
2005. Harvard University Press, 79 Garden Street, Cambridge, 
Massachusetts 02138 USA. 372 pages. US$29.95 Cloth. 


Where to watch birds in Spain. The 100 best sites. By José 
Antonio Montero. March 2006. Lynx Edicions, Montseny, 
8, 08193 Bellaterra, Barcelona, Spain. 362 pages. US$32.50 
Paper. 


Rodents and Lagomorphs of BC. By D. Nagorsen. 2005, 
Royal BC Museum, 675 Belleville Street, Victoria, British 
Columbia V8W 9W2 Canada. 410 pages. 


Annotated Bibliography of the Varanoidea. By M. Bay- 
less. 2007. Krieger Publishing, P.O. Box 9542, Melbourne, 
Florida 32902 USA. 


* Wildlife Demography — Analysis of Sex, Age, and Count 
Data. By John Skalski, Kristin Ryding, and Joshua Mills- 
paugh. 2005. Elsevier Canada,! Goldthorne Avenue, Toronto, 
Ontario M8Z 5S7 Canada. 656 pages. US$69.95 Cloth. 


BOTANY 


Illustrations of Alien Plants of the British Isles. By E. 
Clement, D. Smith and I. Thirlwell. 2005 NHBS Environment 
Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 5XN, United 
Kingdom. £18.75, US$34 Paper. 


Flowers of Crete. By John Fielding and Nicholas Turland. 
2005. Kew Publishing, Royal Botanic Gardens, Kew, Rich- 
mond, Surrey TW9 3AB United Kingdom. 2005. 650 pages. 
£47.00, US$86.00 Cloth. 


Good Wood, Growth, Loss and Renewal. By S. Radose- 
vich. 2005. Oregon State University Press, 102 Adams Hall, 
Corvallis, Oregon 97331 USA. 160 pages. US$18.95. 


New Flowering: 1000 Years of Botanical Art. By S. Sher- 
wood. 2005. NHBS Environment Bookstore, 2-3 Wills Road, 
Totnes, Devon TQ9 5XN United Kingdom. £20 U.S.$36 Paper. 


Orchids of Britain and Ireland — A Field and Site Guide. 
By Anne Harrap and Simon Harrap NHBS Environment 
Bookstore, 2-3 Wills Road, Totnes, Devon TQ9 S5XN United 
Kingdom. £29.99, U.S.$53 Paper. 


Orchids of the British Isles. By M. Foley and S. Clarke. 
2006. Griffin Press. Level 13, 67 Albert Avenue Chatswood, 
New South Wales 2067, in association with the Royal Botanic 
Garden Edinburgh. 320 pages. £45. 


Savannas and Dry Forests: Linking People with Nature. 
By J. Mistry and A. Berardi, 2006. Ashgate Publishing Com- 
pany, Suite 420, 101 Cherry Street, Burlington, Vermont 05401- 
4405 USA. 286 pages. US$114.95. 


The Wild Flower Key — How to identify wild flowers, trees 
and shrubs in Britain and Ireland. By Francis Rose and 
Clare O’ Reilly. March 2006 Frederick Warne Books — Dis- 
tributed by NHBS, 2-3 Wills Road, Totnes, Devon TQ9 SXN 
United Kingdom. 480 pages. £14.99. 


612 


MISCELLANEOUS 


Dry — Life without Water. By E. Masood and D. Schaffer. 
2005. Harvard University Press, 79 Garden Street, Cam- 
bridge, Massachusetts 02138 USA. 174 pages. US$29.95. 


The Emirates: A Natural History. Edited by Peter Hellyer 
and Simon Aspinall. 2005. Trident Press Distributed by NHBS, 
2-3 Wills Road, Totnes, Devon TQ9 5XN, United Kingdom. 
428 pages, £65.00. 


Ecology and Conservation of Steppe-land Birds. Edited 
by Gerard Bota, Manuel B. Morales, Santi Mafiosa, and Jordi 
Camprod6n. 2006. Lynx Edicions, Montseny, 8, 08193 Bel- 
laterra, Barcelona, Spain. 343 pages. US$23.20. 


Nature’s Strongholds — The World’s Great Wildlife 
Reserves. By Laura and William Riley. 2005 Princeton Uni- 
versity Press, 41 William Street, Princeton, New Jersey 08540 
USA. 671 pages. US$49.95 Cloth. 


* A New World of Animals — Early Modern Europeans 
on the Creatures of Iberian America. By Miguel de Astia 
and Roger French. 2005. Ashgate Publishing Ltd., Gower 
House, Croft Road, Aldershot, Hampshire GU11 3HR United 
Kingdom. 276 pages. US$89.95, £50.00 Cloth. 


* Mapper of the Mountains: M. P. Bridgland in the Canadi- 
an Rockies, 1902-1930. By Ian S. MacLaren. 2005. University 
of Alberta Press, Ring House 2, University of Alberta, Edmon- 
ton Alberta T6G 2E1 Canada. 576 pages. $39.95 Can. Paper. 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


The New World Tropics — An Introduction For Natural- 
ists Belize — Costa Rica — Trinidad. By R. Walton and G. 
Dodge. 2005. Milwaukee Public Museum, 800 West Wells 
Street, Milwaukee, Wisconsin 53233 USA. US$24.95. 


A Concise History of Ornithology. By M. Walters. 2005. 
Yale University Press, Box 209040, New Haven, Connecti- 
cutt USA. 256 pages. US$22.00 Paper. 


Return to Wild America — A Yearlong Search for the Con- 
tinent’s Natural Soul. By Scott Weidensaul. 2005. North 
Point Press, Farrar, Straus and Giroux, 19 Union Square West, 
New York, New York 10003 USA. 416 pages. US$26.00 Cloth. 


Saskatchewan — Uncommon Vistas. By J. Conway. 2005. 
University of Alberta Press, Ring House 2, University of 
Alberta, Edmonton, Alberta T6G 2E1 Canada. 156 pages. 
$29.95. 


Taxonomy and Plant Conservation. Edited by Etelka Lead- 
lay and Stephen Jury. 2006. Cambridge University Press, 
Distributed by NHBS, 2-3 Wills Road, Totnes, Devon TQ9 
5XN, United Kingdom. 300 pages. £70.00. 


YOUNG READERS 


Backyard Birding for Kids. By Fran Lee. 2005. Gibbs Smith, 
Publisher, P.O. Box 667 Layton, Utah 84041 USA. 64 pages. 
US$9.95 Paper. 


News and Comment 


Marine Turtle Newsletter (110) 


October 2005. 26 pages: GUEST EDITORIAL: Wild and 
head-started Kemp’s Ridley nesters, eggs, hatchlings, nesting 
beaches and adjoining nearshore waters in Texas should 
receive grater protection — ARTICLES: Strandings of Green 
Turtles along the Saurashtra coast, Gujarat, India — First 
sea turtle tagging programme in Iran — NOTES: Observations 
of Green Turtles in the Lakshadweep Islands, west coast of 
India — Gladiatorial attack on a turtle — Length-width and 
length-weight relationships of Caretta caretta from Italy, 
Mediterranean Sea — Tanzania turtle & dugong conservation 
programme — meeting reports — MTSG UPDATE — BOOK 


Saving the Wild: RENEW 2005 


This glossy colour 20-page booklet is published by RENEW 
(REcovery of Nationally Endangered Wildife) through the 
Canadian Endangered Species Conservation Council is sub- 
titled: An opportunity to Participate in Species Recovery in 
Canada. Contents: Letter from the Canadian Wildlife Directors 
— An introduction to the recovery of wild species in Canada 
— Everyone can contribute to saving the wild — How recov- 
ery works in Canada — Support is crucial in getting the job 
done — Recovery successes — Get invoved with saving the 
wild. Visit website www.speciesatrisk.gc.ca. 


Canadian Species at Risk August 2005 


Issued by the Committee on the Status of Endangered 
Wildlife in Canada (COSEWIC), the list is 64 pages con- 
taining: ABOUT COSEWIC: mandate, membership, definitions 
— SUMMARY TABLES: 500 species designated in the cate- 
gories Extinct (13), Extirpated (22), Endangered (184), 
Threatened (129) and Special Concern (152). In addition, 
187 species have been considered but found to be “Not at 
Risk” (148), or “Data Deficient” (39) (Tables 1-3) — Results 
of May 2005 COSEWIC meeting (Tables 4-5) — Explanation 
of status change symbols for reassessed species — COSEWIC 
ASSESSMENT RESULTS — Table 6: Species assessed and 


REVIEW — ANNOUNCEMENTS — NEWS & LEGAL BRIEFS 
— RECENT PUBLICATIONS. 

The Marine Turtle Newsletter is edited by Brendan J. 
Godley and Annette C. Broderick, Marine Turtle Research 
Group, Centre for Ecology and Conservation, University of 
Exeter in Cornwall, Tremough Campus, Penryn TRIO 9EZ 
United Kingdom; e-mail MTN @seaturtle.org; Fax +44 1392 
263700. Subscriptions and donations towards the production 
of the MTN can be made online at <http://www.seaturtle. 
org/mtn/> or postal mail to Michael Coyne (online Editor) 
Marine Turtle Newsletter, | Southampton Place, Durham, 
North Carolina 27705 USA (e-mail: mcoyne @seaturtle. org). 


For copies, other REWEW publications, or more informa- 
tion contact RENEW @ec.gc.ca; telephone (819) 997-8507; 
fax (819) 994-3684. 

It should be noted that the transforming frog (top centre 
among the eight photographs on the cover) although identified 
on the inside front cover as “Northern Cricket Frog (En- 
dangerd)’, appears to lack Cricket Frog characteristic markings 
and pointed snout but shows more resemblance in head shape 
to a Rana, and may instead be a common Green Frog R. 
clamitans. 


designated Extinct — Table 7: Species placed in a “risk 
category” — Table 8, Species assessed and assigned to the 
“Not at Risk” category — Table 9. Species considered and 
placed in the Data Deficient category — Record of Status 
Re-examinations — Record of Name Changes — Names 
previously used by COSEWIC and synonyms. 

Copies of this publication are available from COSEWIC 
Secretariat, c\o Canadian Wildlife Service, Environment Can- 
ada, Ottawa, Ontario K1 A 0H3. See Web site: www.cosewic. 
gc.ca. COSEWIC Status Reports are available from the Species 
at Risk Act (SARA) Public Registry www.sararegistry.gc.ca. 


613 


The Ottawa Field-Naturalists’ Club Awards for 2004 


IRWIN M. BRobDOo, CHRISTINE HANRAHAN, BEVERLY MCBRIDE, AND ELEANOR ZURBRIGG 


At the Club’s Annual Soirée, held on 30 April 2005, 
at St. Basil’s Church in Ottawa, awards were once again 
given to members, and one non-member, who distin- 
guished themselves by accomplishments in the field 
of natural history and conservation, or by extraordinary 
activity within the Club. There isn’t always a winner 
for every potential award, and this year, once again, 


Charles D. Bird — Honorary Member 


Honorary membership in the Ottawa Field Natura- 
lists’ Club is given to members or non-members in 
recognition of their many years of service to the Club 
or for their contributions to Canadian natural history. 
Our newest honorary member, Dr. Charles D. Bird, is a 
retired cattle farmer from Erskine, Alberta, but qualifies 
for the honour on both counts. On the face of it, that 
might seem a little strange, but, as you will see, this is 
no ordinary cattle farmer. 

Although his family roots were in the central Can- 
adian prairies, Charley Bird was born in Oklahoma. 
He developed a love of the natural world at an early 
age, no doubt due at least in part to his father, Ralph 
Durham Bird, an entomologist/naturalist who was 
teaching at the University of Oklahoma at the time, 
and his mother, Lois, who was a botanist. From the 
age of 10 through his college years, Charley followed 
his father’s interests in insects. He later worked with 
the Northern Insect Survey and then in the field of 
forest entomology. At the University of Manitoba, he 
became fascinated with plant ecology and turned to 
the study of plants. Charley developed a strong inter- 
est in mosses at Oklahoma State University where he 
was doing graduate studies, taking courses from the 
bryologist, Dr. G. K. Ikenberry, eventually earning a 
master’s degree and Ph.D. He returned to Canada in 
1960 for a post-doctoral position at the University of 
Alberta in Edmonton, mainly studying mosses. Two 
years later, he accepted a position in the Biology 
Department at the University of Calgary, turning his 
attention to the bryophytes, lichens and vascular plants 
of Alberta and, indeed, all the prairie provinces. At 
the university, Charley taught courses on all these sub- 
jects as well as biogeography, and he curated the 
growing herbarium as well. His research interests 
were principally in lichen and bryophyte ecology and 
taxonomy, and Charley published dozens of papers 
on mosses, lichens and flowering plants in scholarly 
journals such as The Bryologist and Canadian Journal 
of Botany, and our own Canadian Field-Naturalist. He 


the Anne Hanes Natural History Award was not given. 
On the other hand, a new award was approved by 
Council and awarded for the first time this year: The 
Mary Stuart Education Award. More is said about the 
award in the citation below. The following citations 
for those who received an award were read to the mem- 
bers and guests assembled for the event. 


also became involved in Alberta conservation issues 
and joined local natural history clubs such as the 
Calgary Field Naturalist’s Society and Federation of 
Alberta Naturalists. In 1978, he received the Lauran 
Goulden Award for being the outstanding naturalist in 
Alberta that year. With a strong interest in the local 
flora and fauna, Charley published articles on a variety 
of subjects in the Blue Jay, the Calgary Field Naturalist 
and the Alberta Naturalist,. He has, in fact, published 
close to 300 scholarly articles in all. 

Although Charley decided to take early retirement 
in 1979, trading in his academic tweeds for Alberta 
denim to become a cattle farmer, he never set aside 
his love of the natural world, his commitment to con- 
servation, or his interest in high quality scientific pub- 
lication. He therefore has maintained a close associa- 
tion with his local Buffalo Lake Naturalists Society, 
and he stayed on as an Associate Editor of The Cana- 
dian Field-Naturalist, a job he began in 1974 and con- 
tinues to this day, contributing over 30 years of serv- 
ice to the Ottawa Field-Naturalists’ Club. In the post- 
University years, Charley’s interests returned to insects, 
especially butterflies and moths, and he became a major 
contributor to the beautiful and encyclopedic book, 
Alberta Butterflies, published in 1995. At the present 
time, Charley is working on a catalogue of the micro- 
moths of southern Alberta. 

For his achievements in teaching and research, the 
Alberta Society of Professional Biologists presented 
him with the J. Dewey Soper Award in 2000, also mak- 
ing him an Honorary Life Member of that society. 

Whether he is cataloguing mosses, studying the tax- — 
onomy of a lichen genus, working on the phenology ~ 
of flowers, collecting, mounting and identifying hun- — 
dreds of species of tiny moths, tracing his family’s 
genealogy, or, indeed, keeping records of cattle breed- 
ing, Charley is always meticulous, thorough, imagi- — 
native and perceptive... a naturalist’s naturalist. We — 
are therefore proud to add Charley Bird to our roster — 
of distinguished Honorary Members. 


614 


2005 


CLUB AWARDS FOR 2004 


615 


Barbara Barr — Conservation Award (Non-Member) 


The OFNC Conservation Award — Non-Member is 
given in recognition of an outstanding contribution 
by a non-member in the cause of natural history con- 
servation in the Ottawa Valley, with particular empha- 
sis on activities within the Ottawa District. 

Barbara Barr is a very worthy recipient of this award 
in recognition of her outstanding efforts in speaking on 
behalf of natural area protection in the Ottawa area 
through the Regional and municipal planning process- 
es. Barbara 1s an active, dedicated and highly credible 
representative for the Greenspace Alliance of Canada’s 
Capital as well as for the South Keys/Greenboro 
Community Association. 

The thorough, intelligent work that Barbara does 
reflects her excellent grasp of environmental issues in 
the Ottawa area as well as her grasp of the way the 
City works. She is able to be highly effective, for 
example at meetings of the City of Ottawa’s Planning 
and Environment Committee, by accessing informa- 
tion, preparing and presenting thorough briefs, fol- 
lowing-through on outcomes of presentations, and 
providing a reliable and reputable source of sage 
advice to policy makers on natural area conservation 
issues. In this way, she is a highly effective partici- 
pant during the public consultation phases of land 
development proposals, a voice that the conservation 
community can be proud to have speaking on its 
behalf. Public officials appreciate the value that she 
adds to the public consultation process and have 
invited her to participate on various Public Advisory 
Committees. 

Barbara’s frequent interventions at planning meet- 
ings have served to: 

* sustain the importance of natural area conservation 


in the minds of elected officials, by keeping it a vis- 
ible issue; 

raise the consciousness of Councilors on specific 
issues, such as how destructive country lot subdi- 
visions are to Rural Natural Features; 

re-enforce with City staff and Councilors alike the 
importance of a good Planning Process, and of hon- 
oring the commitments, intent and spirit of the Offi- 
cial Plan with respect to Environmental Features 
and Environmental Impact Statements; 

e demonstrate the value-added of public consultation 
and participation in the planning process (seek win- 
win solutions); and 

influence decisions being made on various land use 
proposals. 

Barbara’s interventions have covered a suite of top- 
ics, ranging from specific conservation issues such as 
protection of the Montfort Woods and Leitrim wet- 
land parcels and the Alta Vista Transportation Corridor, 
to more general topics including proposed rural devel- 
opments for country lot subdivisions, suggestions to 
improve the City’s planning process and documents 
and the Urban Natural Areas Environmental Evaluation 
Study. 

One of Barbara’s important characteristics is her 
optimism and determination to stay with the process. 
With economic and land development activities boom- 
ing in the Ottawa area, it takes solid, reasonable, in- 
formed contributions to influence the process and to 
advocate for Smart Growth. 

Through this award, OFNC members recognize the 
outstanding efforts of Barbara to influence Regional 
and City planning decisions to include natural area 
conservation. 


Ghislaine Rozon — Conservation Award (Members) 


The OFNC Conservation Award is given in recog- 
nition of a club member who has made an outstanding 
contribution toward protecting our natural environ- 
ment. Ghislaine Rozon is a remarkable woman who 
more than fits this criterion. She is a dynamic, articulate, 
passionate defender of our local greenspace, particular- 
ly the Larose Forest. She has worked non-stop trying 
to preserve this area as a conservation zone, free of 
development. 

Her battle for the forest began in the summer of 
2002 when, newly retired from a long teaching career, 
she discovered that a development proposal for Larose 
Forest was being given serious consideration by the 
United Counties of Prescott-Russell (UCPR). Aghast 
at what this would do to the natural environment of 
the forest, she threw herself into the fight. She con- 
tacted the OFNC Conservation Committee for advice, 
but quickly perceived that she would need a thorough 
understanding of municipal, provincial and federal 
politics, including a good knowledge of the Planning 
Act, if she was to argue with elected officials on the 


necessity of conservation. Then she realized she would 
need to educate herself in the areas of forestry, hydrol- 
ogy and ecology. All of this she did with tremendous 
competence and customary thoroughness. As Ghislaine 
immersed herself in learning everything there was to 
know about the political process, including the Prov- 
incial Policy Statement, Environmental Assessments 
and related acts, the boxes of files, papers, articles, 
reports and letters grew. Her home became “Command 
Central” for those who were adamant that the forest 
must be preserved. Through detailed reading of numer- 
ous documents, she exposed inconsistencies and in- 
accuracies in the material produced to support devel- 
opment in the forest. 

She also knew that gaining public support was ex- 
tremely important and rallied both residents and vari- 
ous conservation groups to the cause. “The Friends of 
Larose Forest” which existed prior to the controversy, 
was revived and with Ghislaine, presented a united 
front for a development-free Larose. She also con- 
tacted politicians at all levels, as well as the various 


616 


media, offering guided tours of the proposed devel- 
opment site so that they could see it for themselves 
and make up their minds based on facts, not hearsay. 

Anyone who has dealt with Ghislaine over the last 
few years knows that it is not unusual to receive e- 
mails written and fired off at 2, 3 or 4 in the morning! 
It appeared that she never slept! These detailed mes- 
sages... lucid, well-reasoned, intelligent and thought- 
provoking... were regularly mailed to a growing list 
of supporters. Suggestions on how people could help, 
lists of addresses for the relevant government officials, 
sample letters, as well as succinct synopses of many 
relevant reports, all were supplied by Ghislaine who 
became the acknowledged and reasoned voice for sav- 
ing Larose Forest. 

As a last resort, Ghislaine and several others ap- 
pealed the issue to the OMB. This was not a step light- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


ly undertaken. The cost for hiring a lawyer and expert 
witnesses was substantial and was absorbed by the 
appellants. In the end the decision was handed down 
in favour of the developers. Despite this crushing blow, 
Ghislaine has not given up her fight to save the forest. 
As she has done from day one, she continues to explore 
ways in which Larose Forest can be preserved as a 
conservation area, remaining vigilant and alert to all 
threats to this special place. Brimming with ideas, she 
has plans for a website, printed resource material and 
possibly an interpretive centre ... this lady is amaz- 
ing! If ever there was an example of how to go about 
preparing to protect natural areas, one need look no 
further than Ghislaine for inspiration, which is why the 
Club awards her the 2004 Conservation Award for 
Members. 


Martha Camfield — George McGee Service Award 


The OFNC George McGee Service Award is given 
in recognition of a member who has contributed sig- 
nificantly to the smooth running of the Club over sev- 
eral years. Martha Camfield is a very worthy recipient 
of this award in recognition of her extraordinary serv- 
ice Over Many, many years as a dedicated volunteer 
with the Macoun Field Club. Martha has also readily 
volunteered to assist with numerous plant surveys 
and inventories in the Ottawa area, and has served on 
Council. 

For many years — as long as many of today’s young 
Macouners remember and longer — Martha has been 
a dynamic force with Macoun Field Club and the 
committee that manages it. She has readily volunteered 
in many capacities with the young naturalists, fre- 
quently serving as a leader for meetings, workshops 
and field trips, and also working to find speakers and 
leaders for weekly meetings and field trips. 

Martha’s great “hands on” discovery workshops are 
a particular favourite of the Macouners. Each work- 
shop is uniquely researched and supported by plentiful 
plant materials that Martha has collected for dissection 
or identification — whether it be a variety of roots, 
seeds, wild flowers, leaves, fruits, shrubs, sedges, exot- 
ic plants, and more. 

Martha commands instant respect from the young 
people, who respond to her curiosity and whole-hearted 
enthusiasm to share and impart some of the truly amaz- 
ing nature of her subject. Interesting materials coupled 
with good basic instruction on how to use a hand lens 
or microscope leads to ready involvement of the young 
people so that by the end of a workshop everyone has 
dirty fingernails and a much better idea of how leaves 
and seeds and roots function. At one session on horse- 
tails, she burned the ends of the plant stalks to show 


how the silica inside would form “glass-like beads” on 
the ends. When studying seeds, the air became delight- 
fully filled with cattail fluff! A testimony to Martha’s 
impact is most appropriately captured in the words of 
one of her workshop participants, who said, “I think 
that the most important thing that Martha taught us 
was the wonder that can be in just one seed. Thank 
you Martha! We all had a fantastic time.” 

As a Macoun field trip leader, Martha had a genuine 
interest in what each young person was finding, hoping 
to further spur his or her curiosity and interests. One of 
Martha’s gifts is to help wake people up to the world 
around them by making the outings and information 
imparted relevant to them. 

Another facet of Martha’s long-time volunteerism 
for The Ottawa Field-Naturalists’ Club has been through 
her ready assistance on botanical surveys and inven- 
tories of natural areas around Ottawa, most notably 
with Albert Dugal. Martha’s love of the outdoors, of 
nature and of plants in particular, and her remarkable 
energy, make her a natural at this. Martha has spent 
many days helping Albert Dugal — walking, collect- 
ing and listing plant species as part of inventories or 
surveys of a long list of natural areas, including the 
Leitrim wetlands, Spring Hill Bog, Poor Quarry Woods, 
Canadian Museum of Nature lands in Aylmer, Fern 
Bank Wetland and the woodlands east of the Ottawa 
Hospital complex at Smythe Road. Most recently, she 
has been working on an inventory of old growth species 
in Emerald Woods, and wetlands that are part of the 
historic Leitrim Complex. 

This truly remarkable service over many years 1s 
most deserving of the George McGee Service Award. 
Thank you, Martha. 


2005 


CLUB AWARDS FOR 2004 


617 


Andrea Howard — Mary Stuart Education Award 


Andrea Howard, Education Coordinator of the 
Eastern Ontario Biodiversity Museum (EOBM), has 
been chosen to be the first recipient of the Mary Stuart 
Education Award for her outstanding, innovative ap- 
proaches to teaching a variety of natural history topics. 
This new award has been established for members, 
non-members or organizations in recognition of out- 
standing achievements in the field of natural history 
education in the Ottawa Region. The new Education 
Award was appropriately named in honour of Mary 
Stuart, who was so passionately interested in establish- 
ing a new generation of young naturalists. 

Andrea Howard is a very gifted, knowledgeable 
and effective teacher who has been the education 
coordinator at the EOBM in Kemptville ever since 
the museum came into being over five years ago. In 
that capacity she has developed innovative natural his- 
tory programs presented at the museum and at various 
venues around the North Grenville community and in 
Ottawa. Andrea brings to her position of education 
coordinator a unique background in classroom teach- 
ing, science (B.S. from Queen’s University), drama 
(B.A. from Queen’s University, and active in a local 
theatre group) and environmental advocacy (worked 
for Pollution Probe). 

Andrea has developed her own series of creative, 
exciting and accurate biodiversity modules, collective- 
ly called “Museum in a Suitcase” which she brings 
into various classrooms or classroom-like situations. 


Marilyn Ward — Member of the Year 


The OFNC’s Member of the Year award recognizes 
the member judged to have contributed the most to the 
club in the previous year. Marilyn Ward is the kind of 
member whose efforts keep things humming quietly 
but surely along. In 2004 Marilyn was particularly busy 
making significant contributions on at least three fronts. 

Anyone who has called the club’s phone number 
recently has spoken with Marilyn. The OFNC has a 
phone number, but it doesn’t have an office. Incoming 
calls have to go somewhere! For the last four years, 
they’ve gone to Marilyn or to an answering machine 
in her front hallway. Whether she gets the message 
by machine or in person, she is a welcoming greeter 
who efficiently answers questions or connects callers 
— be they out-of-town visitors, members or prospec- 
tive members — with the person who can answer their 
question. We know her hospitality extends beyond the 
phone too. She even arranges to meet new members 
at a monthly meeting in order to personally welcome 
them and to hand them their membership packages, 
which she also compiles. The club’s e-mail address 
also leads to Marilyn. 

Marilyn is a valuable member of the Excursions and 
Lectures committee. She is conscientious and reliable, 
and she cares about details. As any field trip leader 


These programs include “Secret Life of the Forest’ (the 
biodiversity of decomposition), “Climate Change” (bio- 
diversity and global warming), “Life Cycles & Seasonal 
Adaptations” (biodiversity of survival), “Introduction 
to Taxonomy” (classification of biodiversity) and “Bug- 
watch/Insects” (biodiversity of insects). Andrea skil- 
fully adapts each program to the particular teacher and 
classroom situation by consulting with the teacher in 
question. The literacy program that Andrea created and 
implemented for local high school students having 
reading difficulties is something that the EOBM is par- 
ticularly proud of. The subject matter employed was, 
of course, natural history. 

The need for training teachers has not escaped 
Andrea’s attention either. She has developed and imple- 
mented professional development programs for teach- 
ers, helping them also to explore the possibilities of 
the immediate neighbourhood, especially the school 
grounds. 

Andrea always seeks to learn more by asking appro- 
priate professionals for help and advice and having 
them review her syllabus. She reads constantly to up- 
grade her knowledge and all of this is reflected in the 
timely and lively content and presentations in her class- 
es. She has an excellent rapport with her students and 
their teachers. For all the above reasons and many 
more, we have chosen Andrea Howard for the Educa- 
tion Award. 


will tell you, she is good at recruiting field trip lead- 
ers. She also phones leaders and speakers in advance 
of their commitments to make sure they haven’t for- 
gotten — a service much appreciated by club mem- 
bers. Not only that, she personally shows up at almost 
every field trip and monthly meeting where she helps 
out with logistical details. 

The Fletcher Wildlife Garden gets a lot of traffic, 
with committee and council meetings, volunteers and 
visitors. Somehow, amazingly, it almost always looks 
neat and tidy. Of course all users are expected to clean 
up after themselves, but we all know this sort of thing 
works better in theory than in practice. Marilyn has 
been described as a fiend when it comes to keeping 
the Fletcher centre orderly! She’s often there at least 
twice a week. She also provides suet cakes for the 
Fletcher bird feeding stations and makes regular walks 
around the whole site to see what birds are around. 

Handling the club phone, helping to make sure 
events are successful, keeping the place spic and span: 
we can’t underestimate the importance of these con- 
tributions as part of the public face of the club. Marilyn, 
the club is grateful for your many special contribu- 
tions during 2004. Designating you as Member of the 
Year is our way of saying, “ Thank you!” 


Index to Volume 119 
Compiled by Leslie Cody 


Abies amabilis, 160 
balsamea, 129,255,381,457,570 
lasiocarpa, 186 
Acanthomyops subglaber, 39 
Accipiter cooperii, 379 
striatus, 377 
Acer negundo, 220,438 
Acer pensylvanicum, 129 
Acer rubrum, 17,240,386 
Acer saccharinum, 129 
Acer saccharum, 38,129,332,378,386,457 
Acer spicatum, 457 
Achaearanea ohlerti, 269 
tabulata, 269 
tepidariorum, 269 
Achigan a grande bouche, 363 
a petite bouche, 363 
Achillea millefolium, 248 
millefolium f. rosea, 438 
ptarmica, 438 
Acjaearamea tabulata, 272 
Aconitum sp., 438 
Actaea rubra f. neglecta, 438 
Actitis macularia, 69,379 
Aculepeira carbonarioides, 256 
Adkins-Giese, C.L., and F.J. Cuthbert. Woodpecker Nest Tree 
Characteristics in Upper Midwestern Oak Forests, 367 
Aegithalus caudata, 120 
Aegolius acadicus, 367,578 
funereus, 367,578 
Aegolius acadicus, Caching Behavior by Wintering Northern 
Saw-Whet Owls, 578 
Aesculus hippocastanum, 344 
Agelaius phoeniceus, 69 
Agelenopsis, 95 
utahana, 256 
Agroeca ornata, 266 
Agropyron sericeum, 279 
smithii, 220 
trachycaulum, 78, 279 
Agrostis humilis, 417 
scabra, 394 
thurberiana, 420 
Agyneta sp., 259 
allosubtilis, 259 
amersaxatilis, 259 
dynica, 259 
fabra, 259 
Jacksoni, 259 
olivacea, 259 
simplex, 259 
Aix sponsa, 69,379,548 
Alaska, Attempted Predation of a Child by a Gray Wolf, Canis 
lupus, near Icy Bay, 197 


618 | 


Alaska Beaufort Sea Lagoons, 1999-2002, Long-tailed Duck, 
Clangula hyemalis, Eider, Somateria spp., and Scoter, 
Melanitta spp., Distributions in Central, 181 

Alberta Alfalfa Fields, Body Weights of Adult and Juvenile! 
Northern Pocket Gophers, Thomomys talpoides, in 
Central, 551 

Alberta’s Tardigrade Fauna, Water-bears from the Roc 
Mountains: A First Look at, 586 

Alberta, Travel Rates of Wolves, Canis lupus, in Relation to 
Ungulate Kill Sites in Westcentral, 573 

Alces alces, 43,50,136,168,186,194,197,323,330,381,574 

Alces alces, and Beaver, Castor canadensis, in Algonquin) 
Park, Ontario, 1860-2004, Reconstructing Changes: 
in Abundance of White-tailed Deer, Odocoileus vir- 
ginianus, Moose, 330 

Alces alces, in Central Interior British Columbia, Winter 
Habitat Use by Moose, 186 

Alder, 96,591 
Speckled, 65 

Alewife, 18 

Alfalfa, 175,551 . 

Alfalfa Fields, Body Weights of Adult and Juvenile North: 
ern Pocket Gophers, Thomomys talpoides, in Centra’ 
Alberta, 551 

Alfalfa Fields Inhabited by Northern Pocket Gophers, Tho: 
momys talpoides, Long-tailed Weasel, Mustela fre’ 
nata, Movements and Diggings in, 175 

Alga, Marine Green, 82 

Alga Percursaria percursa from Hypersaline Springs in the 
Middle of the North American Continent, Identifica’ 
tion of a Marine Green, 82 q 

Alisma plantago-aquatica var. americanum, 279 
triviale, 277 

Allan, M., 453 

Allomengea dentisetis, 259 

Alnus spp., 96,386,591 
incana, 389 
incana americana, 96 
incana ssp. rugosa, 65,129,388 
rugosa, 96 

Alopecosa aculeata, 266 

Alopecurus arundinaceus, 438 
geniculatus, 418 
occidentalis, 438 
pratensis, 418 

Alopex lagopus, 120,323 

Alosa pseudoharengus, 21 i) 

Amara aulica, 271 | 

Amaranthus spp., 548 

Amaurobius borealis, 256 | 

Ambloplites rupestris, 21 i 

Ambylpone pallipes, 39 | 

Ambystoma gracile, 292 | 
macrodactylum, 291 


2006 


Ambystoma macrodactylum, in the Oregon Cascade Range, 
Apparent Predation by Gray Jays, Perisoreus canaden- 
sis, on Long-toed Salamanders, 291 
Ameiurus nebulosus, 21,295,360 
Amelanchier alnifolia, 78 
laevis, 240 
lucida, 239 
Amia calva, 21 
Ammodramus leconteii, 69 
Ammodytes americanus, 216 
Ammophila harti, 9 
Amsinckia menziesii, 438 
Anas acuta, 69,548 
americana, 69 
clypeata, 69,548 
crecca, 69,548 
discors, 69,379,548 
platyrhynchos, 69,379,541,546 
rubripes, 69 
strepera, 548 
Anas platyrhynchos, Use of Radio-Telemetry to Test for Inves- 
tigator Effects on Nesting Mallards, 541 
Anax junius, 234 
Andersen, D.E., 208 
Andrena spp., 53,249 
Andrenid, 53 
Andromeda polifolia, 210 
Anemone parviflora, 276 
richardsonii, 276 
Anemone, Plumose, 355 
Angelica lucida, 417 
Angelica, Seacoast, 433 
Anguilla rostrata, 294,351 
Ant Species (Hymenoptera: Formicidae) Richness at the 
Mont St. Hilaire Biosphere Reserve, Québec, The 
Effect of Human Activity on, 38 
Antennaria isolepis, 276 
pulcherrima, 417 
Anthus spragueii, 490 
Antistea brunnea, 259 
Apamea amputatrix, 12 
ophiogramma, 12 
Aphaenogaster rudis, 39 
Aphileta misera, 259 
Aphragmus eschscholtzianus, 417 
Apis mellifera, 53,249 
Aplodinotus grunniens, 21 
Apocynum androsaemifolium, 417 
Arabidopsis salsuginea, 417 
Arabis boivinii, 417 
Aralia nudicaulis, 44 
Araneus corticarius, 256 
diadematus, 255 
displicatus, 257 
groenlandicola, 257 
marmoreus, 257 
nordmanni, 257 
saevus, 257 
solitarius, 257 
trifolium, 257 
washingtoni, 257 
Araniella displicata, 257 
proxima, 257 
Arcella, 403 
arenaria var. grandis, 410 


INDEX TO VOLUME 119 


619 


artocrea, 410 
artocrea ssp. pseudocatinus, 410 
bathystoma, 410 
crenata, 410 
formosa, 403 
leidyana, 410 
megastoma, 410 
rota, 410 
Arcella formosa n. sp.: Two New Species of Testate Rhizo- 
pods (Arcellinida, Protozoa) from Remnant Wetlands 
in Ontario, Canada, Cyclopyxis acmodonta n. sp. and, 
403 
Archilochus colubris, 70,379 
Arctosa alpigena, 266 
emertoni, 267 
insignita, 267 
lama, 267 
quinaria, 267 
raptor, 267 
rubicunda, 267 
virgo, 267 
Arctostaphylos spp., 210 
alpina, 277 
uva-ursi, 240 
Ardea herodias, 69,379 
Argenna obesa, 258 
Argyneta cauta, 259 
decora, 259 
Arnica alpina ssp. attenuata, 289 
angustifolia ssp. attenuata, 277 
chamissonis, 277 
diversifolia, 417 
latifolia, 417 
Arnica, Diverse, 434 
Mountain, 435 
Aronia melanocarpa, 240 
Arrow-grass, Seaside, 279 
Artemisia sp., 220,532 
absinthium, 438 
dracunculus, 438 
furcata, 277 
hyperborea, 289 
tilesii, 277,417 
Asclepias syriaca, 525 
Ash, Mountain, 570 
Asio flammeus, 70,400 
otus, 395 
Asio otus: A Review of North American Banding, Long-eared 
Owls, 395 
Aspen, 175,369,401 
Big-toothed, 371 
Quaking, 370 
Trembling, 65,135,186,220,370,381,442,45 1,574 
Asphodel, Sticky False, 428 
Aspicilia, 78 
caesiocinerea, 78 
Aster brachyactis, 418 
curtus, 245 
hesperius, 439 
lanceolatus, 390 
lanceolatus ssp. hesperius, 439 
puniceus, 276,388 
pygmaeus, 277 
sibiricus, 277 
yukonensis, 418 


620 


Aster, Lindley’s American, 435 
Purple-stemmed, 289 
Rayless, 435 
Western Willow, 439 
White-top,248 
Yukon, 435 
Yukon-American, 435 

aster du Saint-Laurent, 556 

Astragalus alpinus, 417,506 
australis, 277 
eucosmus, 276 
eucosmus f. albinus, 276 
richardsonii, 287 

Athene culicularia, 490 

Athyrium filix-femina var. cyclosorum, 417 

Atkinson, J., Review by, 305 

Attix, L., 455 

Auditor’s Report, 478 

Aulnes rugueux, 129 

Avena sativa, 418 

Aythya collaris, 69 
valisineria, 546 


Babin, T., 453 
Badger, 584 
North American, 442 
Badiou, P.H., 82 
Baeolophus bicolor, 374 
Baggs, E.M., 323 
Bakeapple, 412 
Balaenoptera acutorostrata, 214 
musculus, 214 
physalus, 214 
Balaenoptera acutorostrata, in the Saguenay-St. Lawrence 
National Marine Park, Novel Surface Feeding Tac- 
tics of Minke Whales, 214 
Ballard, W.B., 584 
Baneberry, Red, 438 
Barbotte brune, 360 
Barker, M.E., 58 
Barnes, D.M. Possible Tool Use by Beavers, Castor cana- 
densis, in a Northern Ontario Watershed, 441 
Bartramia longicauda, 69 
Baryphyma kulczynskii, 259 
Bass, Largemouth, 21 
Rock, 21 
Smallmouth, 21,295,353 
White, 21 
Bassaniana utahensis, 270 
Basswood, 369 
Bat, Big Brown, 132 
Hoary, 132 
Little Brown, 444,589 
Bat by American Crows, Corvus brachyrhynchos, Predation 
of a, 443 
Bat, Lasiurus cinereus, Use of a Bridge for Day Roosting by 
the Hoary, 132 
Bathyphantes brevipes, 259 
brevis, 259 
canadensis, 260 
concolor, 260 
crosbyi, 260 
gracilis, 260 
pallidus, 260 
pullatus, 263 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


reprobus, 260 
rufulus, 263 
simillimus, 260 
sp. reprobus, 260 

Bats, Myotis lucifugus, in a Rodent Trap in the Boreal Forest, 
Mortality of Little Brown, 589 

Batt, JH. 355 

Beak-rush, White, 281 

Bear, Black, 50,164,298,323,339,381,574 
Brown, 339,449 
Grizzly, 574 

Bear, Ursus americanus, Ecology on the Northeast Coast of 
Labrador, Black, 164 

Bears, Ursus arctos, from Coastal British Columbia, Obser- 
vations of Autumn Courtship and Breeding in Brown, 
449 

Bearberry, 210 
Alpine, 288 

Beaudoin, A.B., Reviews by, 150,465,596,603 

Beaver, 176,197,330,385,441,591 

Beaver, Castor canadensis, Flooding on a Small Shore Fen 
in Southwestern Quebec, Consequences of, 385 

Beaver, Castor canadensis, in Algonquin Park, Ontario, 1860- 
2004, Reconstructing Changes in Abundance of White- 
tailed Deer, Odocoileus virginianus, Moose, Alces 
alces, and, 330 

Beavers, Castor canadensis, Feeding on Salmon Carcasses: 
Opportunistic Use of a Seasonally Superabundant 
Food Source, 591 

Beavers, Castor canadensis, in a Northern Ontario Water- 
shed, Possible Tool Use by, 441 

Bedstraw, Sweet-scented, 434 

Beech, American, 38,332,378 

Beetle, 271 
Leaf, 88 
Mountain Pine, 186 

Beetles), in North America (Coleoptera: Chrysomelidae, 
Chrysomelinae), New Distribution Records and Bio- 
geography of Calligrapha Species (Leaf, 88 

Bembix americana, 10 
pallidipicta, 9 

Bennett, B.A., 417 

Bennett, K.E., E.M. Baggs, J.R. Finney-Crawley, and M. 
McGrath. Analysis of the Parasites of a Mid-winter 
Population of the Snowshoe Hare, Lepus americanus, 
on Insular Newfoundland During a Cyclical Peak, 
523 

Bennett, R., Review by, 303 

Bentgrass, Alpine, 420 

Berry, C.R: Jr; 219 

Berry, Thorny Buffalo, 220 

Betula sp., 96,186,255,386,413,591 
alleghaniensis, 332,386,457,570 
cordifolia, 570 
glandulosa, 210 
lenta, 96 
papyrifera, 41,96,103,129,381,386,45 1,457,569 
populifolia, 129,240 

Bidens sp., 388,548 
cernua, 394 
connata, 390 

Bidensomela, 89 

Bilberry, Alpine, 412 

Bind-weed, 428 

Birch, 186,255,591 


2006 


Dwarf, 210,413 
Heart-leaved, 570 
Paper, 41,103,371,569 
White, 381,386,451 
Yellow, 332,386,570 

Bistort, 283 

Bitter-cress, 248 

Bittern, American, 69 

Blackbird, Brewer’s, 69 
Red-winged, 69 

Bladderwort, Greater, 137 
Lesser, 434 

Blarina brevicauda, 456,578 

Blarina brevicauda, Apparently Killed by Red Squirrel, 
Tamiasciurus hudsonicus, Short-tailed Shrew, 456 

Blueberries, Vaccinium angustifolium Ait. and V. myrtilloides 
Michx. (Ericacaeae), in the Boreal Forest, Pollination 
and Breeding System of Lowbush, 48 

Blueberry, 44 
Lowbush, 48 

Bluegill, 20 

Bluet, 234 
Familiar, 234 

Blysmopsis rufa, 417 

Blysmus rufus, 422 

Boal, C.W. and D.E. Andersen. Microhabitat Characteristics 
of Lapland Longspur, Calcarius lapponicus, Nests 
at Cape Churchill, Manitoba, 208 

Bobolink, 70 

Boeckner, M.J. and H.C. Proctor. Water-bears from the Rocky 
Mountains: A First Look at Alberta’s Tardigrade 
Fauna, 586 

Bogbean, 433 

Bombus spp., 48 
bifarius, 249 
flavifrons, 249 
ternarius, 56 
terricola, 56 

Bombycilla cedrorum, 70 

Bombylius major, 249 

Bonasa umbellus, 44,71,379 

Bonasia bonasia, 120 

Bondrup-Nielsen, S., 164 

Boreal Dip Net/L’Epuisette Boreale: Newsletter of the Cana- 
dian Amphibian and Reptile Conservation Network/ 
Reseau Canadien de Conservation des Amphibiens 
et des Reptiles, The, 319,473 

Botaurus lentiginosus, 69 

Bouleau blanc, 129,457 
gris, 129 
jaune, 457 

Bouteloua gracilis, 220 

Bowfin, 18 

Boyd, D.K., 446 

Boyd, H., 483 

Boyle, H.F., 6 

Brachinus sp., 16 

Brachymyrmex depilis, 38 

Bradford, R.G., 294 

Brant, Eastern High Arctic, 486 

Branta bernicla hrota, 486 
canadensis, 69,379,483 
canadensis canadensis, 483 
canadensis interior, 483 
hutchinsii, 483 


INDEX TO VOLUME 119 


621 


Branta canadensis, and Cackling Geese, Branta hutchinsii, 
in the Eastern Canadian Arctic, Breeding and Non- 
Breeding Range of Canada, 483 

Branta hutchinsii, in the Eastern Canadian Arctic, Breeding 
and Non-Breeding Range of Canada, Branta cana- 
densis, and Cackling Geese, 483 

Brasenia schreberi, 137 

Braya glabella, 277 
humilis, 277 

British Columbia, Body Size Distribution and Frequency of 
Anthropogenic Injuries of Bluntnose Sixgill Sharks, 
Hexanchus griseus, at Flora Islets, 537 

British Columbia, Insect Visitation to Wildflowers in the 
Endangered Garry Oak, Quercus garryana, Ecosys- 
tem of, 245 

British Columbia, Observations of Autumn Courtship and 
Breeding in Brown Bears, Ursus arctos, from Coastal, 
449 

British Columbia’s Coastal Archipelago, Facts from Faeces: 
Prey Remains in Wolf, Canis lupus, Faeces Revise 
Occurrence Records for Mammals of, 192 

British Columbia, The Distribution and Habitat Selection of 
Introduced Eastern Grey Squirrels, Sciurus carolinen- 
sis, in, 343 

British Columbia, Winter Habitat Use by Moose, Alces alces, 
in Central Interior, 186 

Brodiaea, Howell’s, 248 

Brome, Japanese, 421 

Bromus japonicus, 417 

Brooklime, American, 434 

Broom, Scotch, 245 

Broomrape, Clustered, 434 

Bubo scandiacus, 211 
virginianus, 62,71,395,578 

Bucephala clangula, 69 

Buchanan, L., 453 

Buckbean, 433 

Buckbrush, 220 

Buckwheat, Wild, 428 

Buddle, C.M., 38 

Buffalo, Bigmouth, 18 

Bufo boreas, 291 

Bullfinch, 120 

Bullhead, Brown, 21,295 

Bulrush, 16 
Common Three-Square, 439 
Great, 17 
Hard-stemmed, 427 
Red, 422 

Bumblebee, 48 

Bunchberry, 44,412 

Bunting, Snow, 120 

Bur-reed, 16,137 

Bury, R.B., 291 

Buteo lineatus, 377 
platypterus, 71,377 
regalis, 490 

Butter-and-eggs, 14 

Buttercup, Western, 248 
White Water, 285 

Butterfly, Monarch, 525 


Cactus, Prickly Pear, 220 
Calamagrostis canadensis, 77 
lapponica, 277 


622 


neglecta, 279 

purpurascens, 277 

stricta ssp. stricta, 277 
Calcarius lapponicus, 208 
Calcarius lapponicus, Nests at Cape Churchill, Manitoba, 

Microhabitat Characteristics of Lapland Longspur, 

208 
Calidris alpina, 379 

minutilla, 379 
Calla palustris, 388 
Calliergon stramineum, 388 
Calligrapha, 88 

alni, 89 

alnicola, 89 

amator, 89 

amelia, 89 

androwi, 95 

apicalis, 89 

bidenticola, 91 

californica, 92 

cephalanti, 95 

confluens, 92 

dislocata, 95 

dolosa, 96 

floridana, 95 

fulvipes, 95 

ignota, 89 

incisa, 96 

knabi, 92 

lunata, 96 

multiguttata, 95 

multipunctata, 96 

ostryae, 92 

philadelphica, 93 

pnirsa, 96 

praecelsis, 96 

pruni, 96 

rhoda, 93 

rowena, 89 

scalaris, 93 

sensu Stricta, 89 

serpentina, 95 

sigmoidea, 96 

spiraea, 94 

stolonifera, 96 

suturella, 94 

sylvia, 95 

tiliae, 95 

verrucosa, 95 

vicina, 95 

virginea, 96 

wickhami, 95 
Calligrapha Species (Leaf Beetles), in North America 

(Coleoptera: Chrysomelidae, Chrysomelinae), New 

Distribution Records and Biogeography of, 88 
Callioplus euoplus, 256 

tibialis, 256 
Callitriche hermaphroditica, 277 
Callobius bennetti, 256 
Calophasia lunula, 12 
Caltha natans, 276 

palustris var. arctica, 277 
Camas, Common, 245 
Camassia quamash, 245 
Camomile, Sea-shore, 290 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Campanula rapunculoides, 438 
rotundifolia, 417 
uniflora, 276 

Camponotus nearcticus, 39 
novaeboracensis, 39 
pennsylvanicus, 39 

Campostoma anomalum, 21 

Campylium sp., 389 
polygamum, 388 

Canada? A Specimen from the Arctic Archipelago, North- 
west Territories Links Two Allopatric Species of Alka- 
li Grass, Puccinellia, Addition to the Flora of, 497 

Canada, Branta canadensis, and Cackling Geese, Branta 
hutchinsii, in the Eastern Canadian Arctic, Breeding 
and Non-Breeding Range of, 483 

Canada, Cyclopyxis acmodonta n. sp. and Arcella formosa n. 
sp.: Two New Species of Testate Rhizopods (Arcel- 
linida, Protozoa) from Remnant Wetlands in Ontario, 
403 

Canada, First Record of the Plains Minnow, Hybognathus 
placitus, in, 219 

Canada, Papillate Watermeal, Wolffia brasiliensis, in East- 
ern Ontario: An Addition to the Flora of, 137 

Canada III, Additions and Range Extensions to the Vascular 
Plant Flora of the Continental Northwest Territories 
and Nunavut, 276 

Canadian Arctic, Breeding and Non-Breeding Range of Cana- 
da, Branta canadensis, and Cackling Geese, Branta 
hutchinsii, in the Eastern, 483 

Canadian Field-Naturalist, Advice for Contributors to The, 
158,482,642 

Canadian Field-Naturalist 119(1), Erratum: The, 320 

Canadian Journal: Wildlife Afield, New, 156 

Canadian Species at Risk, 613 

Canadian Waters, First Records of Long-beaked Common 
Dolphins, Delphinus capensis, in, 110 

Canis latrans, 1,134,139,161,200,324,381,574,580,584 
lupus, 1,101,135,192,197,381,442,446,573 
lupus beothucus, 323 
lycaon, 139,330 

Canis latrans, in Labrador, Evidence of Range Expansion 
of Eastern Coyotes, 381 

Canis latrans, in New York, Chronology of Range Expansion 
of the Coyote, 1 

Canis latrans, Record Size Female Coyote, 139 

Canus lupus, Den and Associated Human Activity in the 
Southwestern Yukon Territory, An Ancient Wolf, 135 

Canis lupus, Faeces Revise Occurrence Records for Mam- 
mals of British Columbia’s Coastal Archipelago, Facts 
from Faeces: Prey Remains in Wolf, 192 

Canis lupus, in Relation to Ungulate Kill Sites in Westcen- 
tral Alberta, Travel Rates of Wolves, 573 

Canis lupus, in the Wild, Longevity and Productivity of Three 
Wolves, 446 

Canis lupus, near Icy Bay, Alaska, Attempted Predation of a 
Child by a Gray Wolf, 197 

Canvasback, 546 

Capelin, 215,355 

Capercaillie, 120 

Cappuccino, N., 525 

Carassius auratus, 21 

Carbyn,)S:, 237 

Cardamine sp., 248 
bellidifolia, 276 
pratensis, 276 


2006 


Cardinal, Northern, 71 
Cardinalis cardinalis, 71 
Carduelis flammea, 120 
hornemanni, 120 
pinus, 71 
tristis, 71 
Carex sp., 16,186,388,548,591 
aquatilis, 210 
atherodes, 77 
bebbii, 277,394,417 
brunnescens, 417 
buxbaumii, 417 
canescens, 276,417 
comosa, 137 
cryptolepis, 388 
eburnea, 417 
enanderi, 425 
lasiocarpa, 389,417 
laxa, 417 
lenticularis var. dolia, 417 
lenticularis var. lipocarpa, 417 
livida, 417 
media, 276 
microglochin, 417 
microptera, 417 
nardina, 277,418 
nigricans, 417 
parryana, 418 
phaeocephala, 418 
pseudo-cyperus, 389 
rariflora, 276 
rotundata, 276 
sartwellii, 417 
scirpoidea, 276 
siccata, 417 
subspathacea, 277 
supina ssp. spaniocarpa, 277 
tonsa var. rugosperma, 240 
vaginata, 277 
Caribou, 43,119,168,201,355,381 
Peary, 120 
Woodland, 135,573 
Carorita limnaea, 260 
Carp, Common, 18,546 
Carpiodes cyprinus, 21 
Carpodacus purpureus, 70 
Carya spp., 343 
cordiformis, 369 
ovata, 369 
Cassiope stellariana, 433 
tetragona, 506 
Castanea dentata, 343 
Castilleja miniata, 418 
parviflora, 418 
unalaschcensis, 418 
Castor canadensis, 176,197,330,385,441,591 
Castor canadensis, Feeding on Salmon Carcasses: Oppor- 
tunistic Use of a Seasonally Superabundant Food 
Source, Beavers, 591 
Castor canadensis, Flooding on a Small Shore Fen in South- 
western Quebec, Consequences of Beaver, 385 
Castor canadensis, in Algonquin Park, Ontario, 1860-2004, 
Reconstructing Changes in Abundance of White-tailed 
Deer, Odocoileus virginianus, Moose, Alces alces, 
and Beaver, 330 


INDEX TO VOLUME 119 


623 


Castor canadensis, in a Northern Ontario Watershed, Possible 
Tool Use by Beavers, 441 
Caswell, P., 417 
Catbird, Gray, 70 
Catchfly, Balkan, 439 
Catfish, Channel, 21 
Catharus fuscescens, 70 
guttatus, 70 
ustulatus, 70 
Catling, P.M. Additions to the Flora of the Continental North- 
west Territories from the Great Slave Lake Area, 437 
Catling, P.M. A Potential for the Use of Dragonfly (Odonata) 
Diversity as a Bioindicator of the Efficiency of Sewage 
Lagoons, 233 
Catling, P.M. Identification and Status of the Introduced Black 
Pine, Pinus nigra, and Mugo Pine, Pinus mugo, in 
Ontario, 224 
Catling, P.M., Review by, 597 
Catling, P.M. and S. Carbyn. Invasive Scots Pine, Pinus syl- 
vestris, Replacing Corema, Corema conradii, Heath- 
land in the Annapolis Valley, Nova Scotia, 237 
Catostomus commersoni, 21,221,295,360 
Cattail, 16,72,221,403 
Broad-leaved, 17 
Common, 420 
Cedar, Eastern White, 386 
Northern White, 404 
Western Red, 160,344 
Celastrina echo, 249 
Celery, Wild, 546 
Centromerita bicolor, 260 
Centromerus bicolor, 260 
cornupalpis, 260 
denticulatus, 260 
furcatus, 260 
latidens, 260 
longibulbus, 260 
persolutus, 260 
sylvaticus, 260 
Centropyxis, 410 
impressa, 407 
Cerastium alpinum, 277 
arvense, 248 
beeringianum, 277 
semidecandrum, 248 
Ceraticelus alticeps, 260 
atriceps, 260 
bulbosus, 260 
fissiceps, 260 
laetabilis, 260 
laetus, 260 
minutus, 260 
similis, 260 
Ceratinella brunnea, 260 
ornatula, 260 
parvula, 260 
Ceratinopsis nigriceps, 261 
nigripalpis, 261 
Ceratophyllum demersum, 16,137,548 
Cervus elaphus, 574 
Ceryle alcyon, 69,293,379 
Ceryle alcyon, During Fall Migration, Territorial Behavior in 
Belted Kingfishers, 293 
Cestode, 324 
Cetraria, 506 


624 


Chaenorhinum minus, 438 
Chalcoscirtus alpicola, 268 
Chamaedaphne calyculata, 388 
Chapleau, F., 359 
Chara sp., 16,26 
altaica, 32 
aspera, 33 
buckellii, 32 
canescens, 26 
contraria, 35 
delicatula, 34 
evoluta, 26 
fragilis, 35 
globularis, 34 
hirsuta, 32 
sibirica, 32 
zeylanica, 30 
Chara canescens, Charophytes of Insular Newfoundland II: 
Chara evoluta and, 26 
Chara evoluta and Chara canescens, Charophytes of Insular 
Newfoundland II: 26 
Charadrius montanus, 532 
vociferus, 70,379 
Charadrius montanus, Differential Parental Care by Adult 
Mountain Plovers, 532 
Chatte de l’est, 360 
Chaulk, K., S. Bondrup-Nielsen, and F. Harrington. Black 
Bear, Ursus americanus, Ecology on the Northeast 
Coast of Labrador, 164 
Chen caerulescens, 212,483 
caerulescens caerulescens, 497 
rossi, 483 
Chéne rouge, 129 
Cheniseo sphagnicultor, 261 
Chenopodium spp., 548 
dessicatum, 277 
leptophyllum, 283,437 
Cherry, Black, 371 
Chestnut, American, 343 
Horse, 344 
Chickadee, 121 
Black-capped, 71,367 
Boreal, 70,367 
Chicken, Greater Prairie, 507,515 
Chickweed, Field, 248 
Chimaerid, 539 
Chipmunk, 578 
Eastern, 323 
Chironomidae, 548 
Chlidonias niger, 69,78 
Chou puant, 129 
Christie, K.S. and T.E. Reimchen. Post-Reproductive Pacific 
Salmon, Oncorhynchus spp., as a Major Nutrient Source 
for Large Aggregations of Gulls, Larus spp., 202 
Chrysemys picta, 389 
Chrysosplenium tetrandrum, 277 
Chub, Creek, 295,351 
Lake, 221,351 
Chubbs, T.E. and E.R. Phillips. Evidence of Range Expan- 
sion of Eastern Coyotes, Canis latrans, in Labrador, 
381 
Chytonix sensilis, 12 
Cicuta bulbifera, 388 
Cinquefoil, Bipinnate, 431 
Circus cyaneus, 69,400 


THE CANADIAN FIELD-NATURALIST 


Vols TLS 


Cirsium foliosum, 418 
Cistothorus palustris, 69 
CITES Control List 2005, 319 
Cladina sp., 78,413 
stellaris, 78 
Cladonia, 78 
Cladophora, 84 
Clangula hyemalis, 181 
Clangula hyemalis, Eider, Somateria spp., and Scoter, Melanit- 
ta spp., Distributions in Central Alaska Beaufort Sea 
Lagoons, 1999-2002, Long-tailed Duck, 181 
Clark, H.O. Jr. Aggressive Behaviour Exhibited by a San 
Joaquin Kit Fox, Vulpes macrotis mutica, 134 
Claytonia perfoliata, 248 
Cleavers, 248,439 
Clematis occidentalis, 418 
occidentalis ssp. grosseserrata, 429 
Clematis, Purple, 429 
Clethrionomys spp., 122 
gapperi, 43,123,323,414 
rutilus, 445 
Cloudberry, 210,412 
Club-moss, Mountain, 278 
Sitka, 418 
Clubiona abbotii, 257 
bryantae, 257 
canadensis, 257 
furcata, 257 
Johnsoni, 258 
kastoni, 258 
kulczynskii, 258 
mixta, 258 
moesta, 258 
norvegica, 258 
obesa, 258 
opeongo, 258 
riparia, 258 
trivialis, 258 
Clubrush, Hudson Bay, 427 
Soft-stem, 427 
Clupea harengus, 216 
pallasi, 112 
Coccothraustes vespertinus, 7\ 
Coccyzus erythropthalmus, 71 
Cody, W.J., Reviews by, 150,463 
Cody, W.J. and K.L. Reading. Additions and Range Exten- 
sions to the Vascular Plant Flora of the Continental 
Northwest Territories and Nunavut, Canada III, 276 
Cody, W.J., B.A. Bennett, and P. Caswell. New Records of 
Vascular Plants in the Yukon Territory VII, 417 
Coeloglossum viride ssp. bracteatum, 418 
Colaptes auratus, 70,369,379 
Collinsia clypiella, 262 
palmeni, 262 
parviflora, 248 
Colpodium, 498 
wrightii, 498 
Comandra umbellata ssp. pallida, 418 
Commandra, Pale, 428 
Commorant, Double-crested, 69 
Comptonia peregrina, 9,240 
Consaul, L.L., L.J. Gillespie, and K.I. MacInnes. Addition 
to the Flora of Canada? A Specimen from the Arctic 
Archipelago, Northwest Territories Links Two Allo- 
patric Species of Alkali Grass, Puccinellia, 497 


2006 


Contopus virens, 71 

Cook, F.R., Reviews by, 141,145,148,154 

Cooke, S.J., Review by, 147 

Coontail, 16,137 

Coot, American, 69 

Coras montanus, 256 

Coregonus huntsmani, 294 

Coregonus huntsmani, from the Wild, First Record of Age 
0+ Atlantic Whitefish, 294 

Corema conradii, 237 

Corema, 237 

Corema conradii, Heathland in the Annapolis Valley, Nova 
Scotia, nvasive Scots Pine, Pinus sylvestris, Replac- 
ing Corema, 237 

Corema, Corema conradii, Heathland in the Annapolis Val- 
ley, Nova Scotia, Invasive Scots Pine, Pinus sylvestris, 
Replacing, 237 

Coreopsomela, 89 

Coriarachne versicolor, 270 

Corixidae, 548 

Cormorant, Double-crested, 72 

Cornicularia auranticeps, 266 
cuspidata, 266 
karpinskii, 266 
minuta, 266 
unicornis, 266 
vigilax, 266 

Cornus, 96 
canadensis, 44,412 
racemosa, |7 
sericea 

Corvus brachyrhynchos, 70,291,443 
corax, 70,291,444 

Corvus brachyrhynchos, Predation of a Bat by American 
Crows, 443 

Corylus spp., 369 
americana, 17,96 
cornuta, 457 

Cotton-grass, Slender, 426 

Cottonwood, 17 
Plains, 132 

Cottus sp., 582 

Coturnicops noveboracensis, 69 

Couesius plumbeus, 221,351 

Cougar, 574,580 

Cougar, Puma concolor, in the Yukon, First Confirmation of, 
580 

Couleuvre a collier, 457 
rayée, 458 

Couleuvre a collier, Diadophis punctatus edwardsii, dans \’ est 
du Québec, Extension de |’aire de distribution de la, 
457 

Coydog, 3 

Coyote, 1,134,139, 161,200,574,580,584 
Eastern, 324,381 

Coyote, Canis latrans, in New York, Chronology of Range 
Expansion of the, | 

Coyote, Canis latrans, Record Size Female, 139 

Coyotes, Canis latrans, in Labrador, Evidence of Range 
Expansion of Eastern, 381 

Cranberry, Dry-ground, 210 
Mountain, 288 

Crane, Sandhill, 69 

Crapet-soleil, 360 

Crappie, Black, 21 


INDEX TO VOLUME 119 


625 


White, 21 
Crataegus sp., 17 
Crayfish, 17 
Crematogaster cerasi, 39 
Crepis nana, 277 
Cress, Aleutian, 429 
Saltwater, 429 
Crossbill, Red, 121 
White-winged, 70 
Crotalus, 95 
Crow, American, 70,291,443 
Crowberry, 255 
Black, 210 
Crows, Corvus brachyrhynchos, Predation of a Bat by Amer- 
ican, 443 
Crustulina borealis, 269 
sticta, 269 
Cryphoeca montana, 259 
Cryptogramma crispa var. acrostichoides, 277 
crispa var. sitchensis, 418 
Crystophora cristata, 382 
Ctenium banksi, 269 
boreale, 269 
fuscum, 269 
riparium, 269 
Cuckoo, Black-billed, 71 
Culaea inconstans, 221,360 
Curlew, Long-billed, 490 
Currier (CS 132 
Cutgrass, Rice, 547 
Cuthbert, F.J., 367 
Cyanocitta cristata, 71,377 
Cybaeopsis euopla, 256 
tibialis, 256 
Cybaeota calcarata, 258 
Cyclopyxis, 403 
acmodonta, 403 
arcelloides, 409 
crucistoma, 407 
grospietschi, 407 
impressa, 407 
lobostoma, 408 
stellata, 408 
trilobata, 407 
trilobata var. maxima, 407 
Cyclopyxis acmodonta n. sp. and Arcella formosa n. sp.: 
Two New Species of Testate Rhizopods (Arcellinida, 
Protozoa) from Remnant Wetlands in Ontario, Can- 
ada, 403 
Cyclosa conica, 257 
Cynanchum rossicum, 525 
Cynomys ludovicianus, 532 
Cyperus erythrorhizos, 548 
odoratus, 548 
Cyprinella spiloptera, 21 
Cyprinus carpio, 21,546 
Cypripedium acaule, 240 
Cystopteris fragilis, 276 
montana, 277 
Cytisus scoparius, 245 


Dace, Longnose, 221 
Northern Redbelly, 221 
Pearl, 221 

Dactylis glomerata, 552 


626 


Daisy, Arctic-alpine, 435 
Large-flowered, 435 
Northern, 435 
Subalpine, 435 

Danaus plexippus, 525 

Dandelion, 552 
Rock, 436 

Danthonia spicata, 240,277 

Dard a ventre jaune, 361 

Darimont, C.T., 192 

Darner, Common Green, 234 

Darter, 582 
Blackside, 18 
Iowa, 221 
Johnny, 19 

d’aster du Saint-Laurent, Symphyotrichum laurentianum, aux 
Iles-de-la-Madeleine, Protocole de suivi des popula- 
tions, 556 

Davies, C., 453 

Davies, J.C., 64 

Davis, H., 296 

Death-camas, Meadow, 248 

Deer, Mule, 296,574,580 
White-tailed, 101,330,381,573,580,591 

Deer, Odocoilus hemionus, and Other Mammals, Passage 
Through a Small Drainage Culvert by Mule, 296 

Deer, Odocoileus virginianus, Moose, Alces alces, and Bea- 
ver, Castor canadensis, in Algonquin Park, Ontario, 
1860-2004, Reconstructing Changes in Abundance 
of White-tailed, 330 

de Lafontaine, G. Protocole de suivi des populations d’ aster 
du Saint-Laurent, Symphyotrichum laurentianum, aux 
Iles-de-la-Madeleine, 556 

Delphinapterus leucas, 214 

Delphinus bairdii, 110 
capensis, 110 
delphis, 110 
delphis bairdii, 110 

Delphinus capensis, in Canadian Waters, First Records of 
Long-beaked Common Dolphins, 110 

Dendroctonus ponderosae, 186 

Dendroica caerulescens, 70 
coronata, 70 
magnolia, 70 
palmarum, 69 
pensylvanica, 71 
petechia, 70 
tigrina, 71 
virens, 70 

Dendryphantes nigromaculatus, 268 

Dermacentor albipictus, 330 

Dermatocarpon, 78 
reticulatum, 78 

Deschampsia flexuosa, 239 

Descurainia sophioides, 277 

Desmognathus fuscus, 105 

Desmognathus fuscus, sur la rive nord du fleuve Saint-Lau- 
rent, au Québec, Premieres mentions et répartition de 
la Salamandre sombre du Nord, 105 

de Solla, S.R., 58 

Desroches, J.-F., 129 

Desroches, J.-F. et B. Roussel. Extension de l’aire de distri- 
bution de la Couleuvre a collier, Diadophis puncta- 
tus edwardsii, dans lest du Québec, 457 

Desroches, J.-F. et D. Pouliot. Premieres mentions et répar- 
tition de la Salamandre sombre du Nord, Desmog- 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


nathus fuscus, sur la rive nord du fleuve Saint-Laurent, 
au Québec, 105 
Devine, A. and D.G. Smith. Caching Behavior by Wintering 
Northern Saw-Whet Owls, Aegolius acadicus, 578 
Devries, J.H., 541 
Diadophis punctatus edwardsii, 457 
Diadophis punctatus edwardsii, dans \ est du Québec, Exten- 
sion de l’aire de distribution de la Couleuvre 4a col- 
lier, 457 
Dianthus barbatus, 438 
Dichanthelium acuminatum ssp. fasciculatum, 240 
depauperatum, 240 
Dicrocoelium dentriticum, 324 
Dicrostonyx groenlandicus, 124 
Dictyna alaskae, 258 
annulipes, 258 
arundinacea, 258 
bostoniensis, 258 
brevitarsus, 258 
minuta, 258 
phylax, 258 
Dicymbium elongatum, 261 
nigrum, 261 
Dietrichia hesperia, 261 
Difflugia lobostoma var. impressa, 407 
Dimelaena, 78 
oreina, 78 
Dinsmore, S.J. and F.L. Knopf. Differential Parental Care by 
Adult Mountain Plovers, Charadrius montanus, 532 
Diphasiastrum sitchense, 417 
Diplocentria bidentata, 261 
corynetes, 264 
rectangulata, 261 
Diplocephalus cristatus, 261 
cuneatus, 261 
subrostratus, 261 
Diplodia, 226 
Dipoena nigra, 269 
Dismodicus alticeps, 261 
bifrons decemoculatus, 261 
decemoculatus, 261 
Dock, Golden, 429 
Dodecatheon hendersonii, 248 
Dog, Coyote-domestic, 3 
Dogbane, Spreading, 433 
Dogwood, 17,96 
Dolichoderus plagiatus, 39 
Dolichonyx oryzivorus, 70 
Dolomedes, 255 
fulviatronotatus, 255 
scapularis, 255 
striatus, 255 
triton, 255 
vittatus, 255 
Dolphin, Bottlenose, 217 
Dusky, 217 
Long-beaked Common, 110 
Short-beaked Common, 110 
Dolphins, Delphinus capensis, in Canadian Waters, First 
Records of Long-beaked Common, 110 
Dondale, C.D., 254 
Donovan, D., 453 
Dorosoma cepedianum, 21 
Douglas-fir, 344 
Douglasia ochotensis, 418 
Dove, Mourning, 71,379 


2006 


Draba albertina, 418 
cinerea, 418 
crassifolia, 418 
fladnizensis, 276 
Juvenalis, 277 
lactea, 286 
lonchocarpa var. vestita, 418 
longipes, 285 
nivalis, 276,418 
oligosperma, 418 
scotteri, 418 
stenoloba, 418 
stenopetala, 418 
wahlenbergii, 276 
Draba, Alaska, 430 
Few-seeded, 430 
Slender, 429 
Snow, 430 
Dragonfly (Odonata) Diversity as a Bioindicator of the Effi- 
ciency of Sewage Lagoons, A Potential for the Use 
of, 233 
Drapetisca alteranda, 261 
Drassodes neglectus, 258 
Drepanocladus, 78 
Drosera anglica, 418 
rotundifolia, 388 
Drum, Freshwater, 21 
Drummond, D.R., 135 
Dryas, 506 
crenulata, 418 
hookeriana, 418 
integrifolia, 210 
integrifolia ssp. crenulata, 418 
octopetala ssp. hookeriana, 418 
Dryocopus pileatus, 70,367,379 
Dryopteris fragrans, 276 
spinulosa, 570 
Duck, 78,546 
American Black, 69 
Long-tailed, 181 
Mallard, 69 
Ring-necked, 69 
Wood, 69,379,547 
Duck, Clangula hyemalis, Eider, Somateria spp., and Scoter, 
Melanitta spp., Distributions in Central Alaska 
Beaufort Sea Lagoons, 1999-2002, Long-tailed, 181 
Ducks During Fall Migration in a Prairie Pothole System, 
Heron Lake, Minnesota, Food Habits of Dabbling, 
546 
Dulichium arundinaceum, 390 
Dumetella carolinensis, 70 
Dunbrack, R. and R. Zielinski. Body Size Distribution and 
Frequency of Anthropogenic Injuries of Bluntnose 
Sixgill Sharks, Hexanchus griseus, at Flora Islets, 
British Columbia, 537 
Dunlin, 379 
Dwarf-Primrose, Arctic-Montane, 433 


Echiniscus spitsbergensis, 587 
Echinochloa crusgalli, 548 
Editor’s Report for Volume 118 (2004), 321 
Eel, American, 294,351 
Eelgrass, 16,28 
Eider, 181 

Common, 182 

King, 182 


INDEX TO VOLUME 119 


627 


Eider, Somateria spp., and Scoter, Melanitta spp., Distribu- 
tions in Central Alaska Beaufort Sea Lagoons, 1999- 
2002, Long-tailed Duck, Clangula hyemalis, 181 
Eidmanella pallida, 267 
Eleocharis sp., 394,548 
erythropoda, 439 
obtusa, 394 
Elk, 574 
Elle, E., 245 
Elm, 369 
American, 96,370 
Slippery, 370 
Elymus canadensis, 277 
junceus, 422 
macrourus, 277 
trachycaulus ssp. violaceus, 277 
Emberiza citrinella, 120 
Emblyna annulipes, 258 
phylax, 258 
Emery, R.B., 541 
Empetrum, 255 
nigrum, 210 
Empidonax alnorum, 70 
flaviventris, 69 
minimus, 70 
tyrannus, 70 
Emydoidea blandingi, 389 
Enallagma, 234 
civile, 234 
cyathigerum, 234 
Encalypta sp., 80 
Enoplognatha caricis, 269 
intrepida, 269 
marmorata, 269 
ovata, 269 
Entelecara abrupta, 266 
exigua, 264 
sombra, 261 
Enteromorpha, 84 
Eperigone bryantae, 261 
contorta, 261 
entomologica, 261 
index, 261 
maculata, 261 
trilobata, 261 
undulata, 261 
Epilobium ciliatum, 394 
coloratum, 394 
davuricum, 276 
hornemannii ssp. hornemannii, 418 
latifolium f. albiflorum, 276 
palustre, 277 
Epinette, 457 
blanche, 458 
noire, 457 
Epinoche A cing épines, 360 
a trois épines, 360 
Epitheca cynosura, 234 
Eptesicus fuscus, 132 
Equisetum palustre, 276 
Equus cabalus, 574 
Erable a épis, 457 
a sucre, 129,457 
argenté, 129 
de Pennsylvanie, 129 
Eremophila alpestris, 71 


628 


Erethizon dorsatum, 125,159,177,453 
Erethizon dorsatum, in Second-Growth Conifer Forests, The 
Influence of Thermal Protection on Winter Den Selec- 
tion by Porcupines, 159 
Eridantes utibilis, 261 
Erigeron grandiflorus ssp. arcticus, 418 
humilis, 418 
peregrinus ssp. peregrinus, 418 
Philadelphicus, 394 
pumilus, 418 
uniflorus ssp. eriocephalus, 277,418 
yukonensis, 418 
Erigone aletris, 261 
atra, 261 
blaesa, 261 
dentigera, 261 
dentipalpis, 255 
ephala, 261 
mentasta, 262 
whymperi, 262 
Eriophorum angustifolium, 277 
callitrix, 277 
gracile, 418 
triste, 277 
vaginatum, 277 
viridi-carinatum, 276 
Ero canionis, 267 
Erskine, A.J., Review by, 299 
Erynnis propertius, 249 
Erysimum pallasii, 276 
Erythemis simplicicollis, 234 
Erythronium oregonum, 248 
Esox lucius, 21,359 
masquinongy, 364 
Esox lucius, dans le lac Ramsay, Parc de la Gatineau, Québec, 
et impact sur l’ichtyofaune, Charactéristiques d’une 
population introduit du Grand brochet, 359 
Estrandia grandaeva, 262 
Etheostoma sp., 582 
exile, 221,361 
nigrum, 21 
Eulaira concava, 259 
microtarsus, 262 
Eupatorium maculatum, 394 
perfoliatum, 389 
Euphagus cyanocephalus, 69 
Euphrasia stricta, 390 
Eurycea bislineata, 106 
Euthamia graminifolia, 388 
graminifolia var. graminifolia, 290 
Eutrema edwardsii, 277 
Euxoa incallida, 12 
scandens, \2 
Evarcha hoyi, 268 
Evers, D., 455 
Ewins, P.J., 58 


Fagus grandifolia, 38,129,332,378 

Falco columbarius, 71,379 
peregrinus, 70 
sparverius, 71,379 

Falcon, Peregrine, 70 

Farnell, R., P.G. Hare, and D.R. Drummond. An Ancient Wolf, 
Canus lupus, Den and Associated Human Activity in 
the Southwestern Yukon Territory, 135 

Felis concolor, 574 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Fener, H.M., J.R. Ginsberg, E.W. Sanderson, and M.E. Gomp- 
per. Chronology of Range Expansion of the Coyote, 
Canis latrans, in New York, 1 
Fern, Fragile, 278 
Fragrant Cliff, 278 
Holly, 419 
Lady, 419 
Mountain Bladder, 278 
Sitka Parsley, 419 
Sweet, 9 
Wood, 570 
Fescue, Alaska, 421 
Alpine, 421 
Little, 421 
Festuca brachyphylla, 276,418 
brevissima, 418 
filiformis, 240 
minutiflora, 418 
richardsonii, 277 
rubra ssp. richardsonii, 279 
Finch, Purple, 70 
Finney-Crawley, J.R., 323 
Fir, Amabilis, 160 
Balsam, 255,381,570 
Subalpine, 186 
Fisher, 120,177,194 
Fivefinger, Marsh, 287 
Fleabane, Shaggy, 435 
Yukon, 435 
Flicker, Northern, 70,369,379 
Floricomus rostratus, 262 
Flycatcher, Alder, 70 
Great-crested, 367 
Least, 70 
Yellow-bellied, 69 
Fontaine, A.J., 483 
Ford, J.K.B. First Records of Long-beaked Common Dol- 
phins, Delphinus capensis, in Canadian Waters, 110 
Forget-me-not, Common, 248 
Forktail, Eastern, 234 
Formica aserva, 40 
glacialis, 39 
lasioides, 39 
neogagates, 40 
nepticula, 40 
nitidiventris, 40 
subaenescens, 40 
subsericea, 40 
42™4 North American Moose Conference, 12-16, 2006 Bad- 
deck, Nova Scotia, Canada, 320 
Fowlie, A.R., 582 
Fox, Arctic, 120,323 
Red, 120,323 
San Joaquin Kit, 134 
Swift, 134,584 
Fox, Vulpes macrotis mutica, Aggressive Behaviour Exhibited 
by a San Joaquin Kit, 134 
Fox, Vulpes velox, Den Located Next to a Railroad Track in 
Northwestern Texas, Swift, 584 
Foxtail, Meadow, 421 
Fragaria virginiana, 394 
Francoeur, A., 38 
Fraxinus nigra, 457 
Freeling, S.E., 219 
Fréne noir, 457 
Fritillaria affinis, 248 


2006 


Frog, Boreal Chorus, 291 
Columbia Spotted, 291 
Northern Leopard, 389 
Pickerel, 578 
Froglog: Newsletter of the Declining Amphibian Populations 
Task Force, 156 
Fulica americana, 69 
Fundulus diaphanus, 21,28,295 


Gadwall, 547 

Galium aparine, 248,439 
tinctorium, 394 
trifidum, 276 
triflorum, 418 

Gallant, J.J., 355 

Gallinago delicata, 69 

Gambelia, 95 

Gar, Longnose, 21 

Gasterosteus aculeatus, 28,360 

Gaultheria hispidula,44 
procumbens, 240 

Gavia immer, 455 

Gavia immer, Nest Attendance Patterns Recorded by Remote 
Video Camera, Common Loon, 455 

Gawn, M., Review by, 604 

Gaylussaccia baccata, 240 

Geese, Branta hutchinsii, in the Eastern Canadian Arctic, 
Breeding and Non-Breeding Range of Canada, Bran- 
ta canadensis, and Cackling, 483 

Gentiana affinis, 277 

Geomys spp. 175 

Geopyxella sp., 408 

Geothlypis trichas, 69 

Geranium erianthum, 418 
molle, 248 

Geranium, Dovefoot, 248 
Northern, 432 

Ghelna canadensis, 268 

Gilbert, B.K., 449 

Gillespie, L.J., 497 

Ginsberg, J.R., 1 

Glaucomys sabrinicus, 344 
sabrinus, 591 
volans, 374 

Gleason, J.S., R.A. Hoffman, and J.M. Wendland. Beavers, 
Castor canadensis, Feeding on Salmon Carcasses: 
Opportunistic Use of a Seasonally Superabundant 
Food Source, 591 

Glenostictia, 6 

Glycine max, 547 

Glyphesis scopulifer, 262 

Gnaphosa brumalis, 258 
microps, 258 
muscorum, 258 
parvula, 258 

Gnathonaroides pedalis, 262 

Goat, Mountain, 197,574 

Goby, Round, 18,582 

Goby, Neogobius melanostomus, in the St. Lawrence River 
at Cornwall, Ontario, First Occurrence of the Round, 
582 

Gold, Spring, 248 

Goldeneye, Common, 69 

Goldfinch, American, 71 

Goldfish, 18 


INDEX TO VOLUME 119 


629 


Gomez-Zurita, J. New Distribution Records and Biogeography 
of Calligrapha Species (Leaf Beetles), in North Amer- 
ica (Coleoptera: Chrysomelidae, Chrysomelinae), 88 
Gompper, M.E., | 
Gonatium crassipalpum, 262 
Gonzales, E.K. The Distribution and Habitat Selection of 
Introduced Eastern Grey Squirrels, Sciurus carolinen- 
sis, in British Columbia, 343 
Goodale, W., L. Attix, and D. Evers. Common Loon, Gavia 
immer, Nest Attendance Patterns Recorded by Remote 
Video Camera, 455 
Goodwin, C.E., Reviews by, 144,600 
Goose, Cackling, 483 
Canada, 69,379,483,549 
Lesser Snow, 497 
Ross’s, 483 
Snow, 212,483 
Gooseberry, 369 
Goosefoot, Narrow-leaved, 283,438 
Gopher, Northern Pocket, 175,551 
Valley Pocket, 551 
Gophers, Thomomys talpoides, in Central Alberta Alfalfa 
Fields, Body Weights of Adult and Juvenile North- 
ern Pocket, 551 
Gophers, Thomomys talpoides, Long-tailed Weasel, Mustela 
frenata, Movements and Diggings in Alfalfa Fields 
Inhabited by Northern Pocket, 175 
Gosse, J.W. and B.J. Hearn. Seasonal Diets of Newfound- 
land Martens, Martes americana atrata, 43 
Grackle, Common, 70 
Gramma, Blue 220 
Grammonota angusta, 262 
capitata, 262 
gigas, 262 
maritima, 262 
pictilis, 262 
vittata, 262 
Grand brochet, 359 
Grand brochet, Esox lucius, dans le lac Ramsay, Parc de la 
Gatineau, Québec, et impact sur l’ichtyofaune, Cha- 
ractéristiques d’une population introduit du, 359 
Graphicallo, 89 
Grasby, S.E., 82 
Grass, Alkali, 497 
Alkali Cord, 280 
Alpine Blue, 280 
American Lyme, 421 
Common Panic, 439 
Ice, 422 
Kentucky Blue, 245 
Needle-and-thread, 220 
Orchard, 552 
Reed Canary, 422 
Turtle, 114 
Western Wheat, 220 
Widgeon, 114 
Grass, Puccinellia, Addition to the Flora of Canada? A Speci- 
men from the Arctic Archipelago, Northwest Terri- 
tories Links Two Allopatric Species of Alkali, 497 
Grass-of-Parnassus, Fringed, 431 
Greasewood, 220 
Grebe, Horned, 69 
Pied-billed, 69 
Grosbeak, Evening, 71 
Pine, 121 


630 


Rose-breasted, 71 
Grouse, Black, 120 
Hazel, 120 
Northern Sharp-tailed, 507,515 
Prairie, 507,515 
Prairie Sharp-tailed, 507,515 
Ruffed, 44,71,379 
Sharp-tailed, 507 
Willow, 120 
Grouse”, Tympanuchus cupido X phasianellus, Hybridization 
on Manitoulin Island, Ontario, “Prairie, 507 
Grouse”, Tympanuchus cupido X phasianellus, of Manitoulin 
Island, Ontario, Plumage and Internal Morphology 
of the “Prairie, 515 
Grus canadensis, 69 
Gull, 202 
Bonaparte’s, 203 
California, 203 
Glaucous-winged, 203 
Mew, 203 
Ring-billed, 69 
Thayer’s, 203 
Gulls, Larus spp., Post-Reproductive Pacific Salmon, Onco- 
rhynchus spp., as a Major Nutrient Source for Large 
Aggregations of, 202 
Gulo gulo, 120 
Gyrinophilus porphyriticus, 106 


Habenaria tridentata, 388 

Hahnia cinerea, 259 
glacialis, 259 

Hake, Pacific, 112 

Halenia deflexa, 438 

Halichoerus grypus, 355 

Halictus sp., 249 

Halorates alascensis, 262 
holmgrenii, 262 
palmeni, 262 
pertinens, 262 

Hamas, M.J. Territorial Behavior in Belted Kingfishers, 
Ceryle alcyon, During Fall Migration, 293 

Hammer, U.T., 114 

Haplodrassus hiemalis, 258 
signifer, 258 

Hare, Arctic, 323 
European, 122 
Snowshoe, 43,121,197,323,395,569 

Hare, Lepus americanus, on Insular Newfoundland During 
a Cyclical Peak, Analysis of the Parasites of a Mid- 
winter Population of the Snowshoe, 323 

Hare, P.G.,. 135 

Harebell, 434 

Hares, Lepus americanus, on Kent Island, New Brunswick, 
Tree Recruitment Limitation by Introduced Snow- 
shoe, 569 

Harrier, Northern, 69,400 

Harrimanella stellariana, 418 

Harrington, F., 164 

Harvey-Clark, C.J., J.J. Gallant, and J.H. Batt. Vision an 
its Relationship to Novel Behaviour in St. Lawrence 
River Greenland Sharks, Somniosus microcephalus, 
oad 

Hasselman, D.J., P. Longue, and R.G. Bradford. First Record 
of Age 0+ Atlantic Whitefish, Coregonus huntsmani, 
from the Wild, 294 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Hawk, Broad-winged, 71,377 
Cooper’s, 379 
Ferruginous, 490 
Red-shouldered, 377 
Sharp-shinned, 377 

Hawkweed, Slender, 435 

Hawthorn, 17 

Hazel, 369 

Hazelnut, 17 
American, 96 

Hearn, B.J., 43 

Hedysarum alpinum, 277 
boreale ssp. mackenzii, 277 
mackenzii, 287 

Helophora insignis, 262 
ontariensis, 259 

Hemidactylium scutatum, 129 

Hemidactylium scutatum, a Québec, Québec: limite nord-est 
de l’espéce sur la rive nord du fleuve Saint-Laurent, 
Découverte de la Salamandre a quatre orteils, 129 

Hemlock, 38,332 
Eastern, 386 
Mountain, 291 
Western, 160,197,344 

Hen, Heath, 516 

Hendricks, P., J. Johnson, S. Lenard, and C. Currier. Use of a 
Bridge for Day Roosting by the Hoary Bat, Lasiurus 
cinereus, 132 

Heron, Great Blue, 69,379 

Herring, 203 
Pacific, 112,216 

Hétre a grandes feuilles, 129 

Hexanchus griseus, 537 

Hexanchus griseus, at Flora Islets, British Columbia, Body 
Size Distribution and Frequency of Anthropogenic 
Injuries of Bluntnose Sixgill Sharks, 537 

Hickey, M.B.C. and A.R. Fowlie. First Occurrence of the 
Round Goby, Neogobius melanostomus, in the 
St. Lawrence River at Cornwall, Ontario, 582 

Hickory, 343,369 

Hieracium gracile, 418 
pilosella, 240 

Hierochloe hirta ssp. arctica, 277 
odorata, 280 

Hilaira algida, 265 
aquilonia, 262 
canaliculata, 262 
dubia, 264 
herniosa, 262 
mentasta, 262 

Hippuris vulgaris, 28 

Hirundo rustica, 70 

Hoffman, R.A., 591 

Holyan, J., D.K. Boyd, C.M. Mack, and D.H. Pletscher. 
Longevity and Productivity of Three Wolves, Canis 
lupus, in the Wild, 446 

Honeybee, 250 

Hood, D.J. and R.F. Stocek. The Fathead Minnow, Pimephales 
promelas, in New Brunswick, 351 

Hordeum vulgare, 438 

Horse, 574 

Horsetail, Marsh, 278 

Houston, C.S. Long-eared Owls, Asio otus: A Review of 
North American Banding, 395 

Houston, C.S., Reviews by, 595,596,598,606,607,608 


2006 


Howerter, D.W., 541 
Hudsonia ericoides, 240 
Huettmann, F., Reviews by, 142,151,302,313,314,315,467,601 
Hulteniella integrifolia, 506 
Hummingbird, Ruby-throated, 70,379 
Hybauchenidium gibbosum, 262 
Hybocoptus denticulatus, 262 
Hybognathus spp., 219 

argyritis, 219 

hankinsoni, 219,360 

nuchalis, 219 

placitus, 219 

regius, 219 
Hybognathus placitus, in Canada, First Record of the Plains 

Minnow, 219 
Hydatigera taeniaeformis, 324 
Hydrocotyle americana, 394 
Hydrolagus colliei, 539 
Hylocichla mustelina, 70 
Hylocomium splendens, 404 
Hypericum mutilum, 394 

perforatum, 14,240 
Hypomma marxii, 262 
Hypsibius, 586 

cf. convergens, 587 
Hypsosinga pygmaea, 257 

rubens, 257 
Hyptiotes gertschi, 270 


Ictalurus punctatus, 21 
Icterus galbula, 70 
Ictiobus cyprinellus, 21 
If du Canada, 457 
Impatiens capensis, 394 
Tris setosa ssp. interior, 418 
versicolor, 388 
Ischnura verticalis, 234 
Islandiana alata, 262 
flaveola, 262 
holmi, 262 
princeps, 263 
Isohypsibius cf. tuberculatus, 587 
Iviella sp., 258 


Jacob’s Ladder, Showy, 433 

Jaeger, Parasitic, 211 

Jay, Blue, 71,377 
Gray, 70,101,291 

Jays, Perisoreus canadensis: Adaptations to Boreal Winters, 
Weight-carrying Ability and Caching Behavior of 
Gray, 101 

Jays, Perisoreus canadensis, on Long-toed Salamanders, 
Ambystoma macrodactylum, in the Oregon Cascade 
Range, Apparent Predation by Gray, 291 

John, R., Reviews by, 141,146,149,299,462,594,599 

Johnson, J., 132 

Johnson, S.R., 181 

Joynt, B.L., 541 

Juglans nigra, 348 

Junco, 578 

Juncus albescens, 282 
arcticus, 276 
brevicaudatus, 388 
bufonius, 418 
canadensis, 394 


INDEX TO VOLUME 119 


631 


castaneus ssp. castaneus, 276 
filiformis, 418 
stygius ssp. americanus, 277 
triglumis ssp. albescens, 277 

Jung, T.S. and B.G. Slough. Mortality of Little Brown Bats, 
Myotis lucifugus, in a Rodent Trap in the Boreal For- 
est, 589 

Jung, T.S. and K.S. O’Donovan. Mortality of Deer Mice, 
Peromyscus maniculatus, in Wire Mesh Live-Traps: 
A Cautionary Note, 445 

Jung, T.S. and P.J. Merchant. First Confirmation of Cougar, 
Puma concolor, in the Yukon, 580 

Jung, T.S., K.S. O’Donovan, and T. Powell. Long-distance 
Movement of a Dispersing Deer Mouse, Peromyscus 
maniculatus, in the Boreal Forest, 451 

Juniper, 78 
Ground, 278 

Juniperus, 78 
communis, 240,277 


Kaestneria pullata, 263 
rufula, 263 

Kalmia angustifolia, 240,381 

Kariz, R.M., 186 

Kelly, E., Review by, 311 

Kestrel, American, 71,379 

Kevan, P.G., 48 

Killdeer, 70,379 

Killifish, Banded, 18,28,294 

Kingbird, Eastern, 70 

Kingfisher, Belted, 69,293,379 

Kingfishers, Ceryle alcyon, During Fall Migration, Territorial 
Behavior in Belted, 293 

Kinglet, Golden-crowned, 579 
Ruby-crowned, 70 

Knopf, F.L., 532 

Knotweed, Striate, 428 

Krawchuk, A., K.W. Larsen, R.D. Weir, and H. Davis. Pas- 
sage Through a Small Drainage Culvert by Mule Deer, 
Odocoilus hemionus, and Other Mammals, 296 

Kuker, K.J., J.A. Thomson, and U. Tscherter. Novel Surface 
Feeding Tactics of Minke Whales, Balaenoptera 
acutorostrata, in the Saguenay-St. Lawrence National 
Marine Park, 214 

Kurczewski, FE. and H.F. Boyle. Nesting Behavior, Ecology, 
Seasonal and Geographic Distribution of the Sand 
Wasp, Stictiella emarginata (Hymenoptera: Spheci- 
dae)*, 6 

Kuzyk, G.W., C. Rohner, and F.K.A. Schmiegelow. Travel 
Rates of Wolves, Canis lupus, in Relation to Ungulate 
Kill Sites in Westcentral Alberta, 573 


Labidesthes sicculus, 21 

Labrador Tea, 288,381,412 
Dwarf, 210 

Labrador, A Range Extension for the Rock Vole, Microtus 
chrotorrhinus, in, 412 

Labrador, Black Bear, Ursus americanus, Ecology on the 
Northeast Coast of, 164 

Labrador, Evidence of Range Expansion of Eastern Coyotes, 
Canis latrans, in, 381 

Lacinipolia vicina, 12 

Lady’s-thumb, 428 

Lady’s-smock, 285 

Lagenorhynchus obscurus, 217 


632 


Lagopus lagopus, 44,120 
Lagotis glauca, 277 
glauca ssp. minor, 288 
Lambdina fiscellaria, 324 
Lance, Sand, 216 
Lanius sp., 102,578 
ludovicianus, 490 
Lansing, S.W. A Range Extension for the Rock Vole, Microtus 
chrotorrhinus, in Labrador, 412 
Lapierre, K. and Coad, B.W, Review by, 301 
Larinioides cornutus, 257 
patagiatus, 257 
sclopetarius, 257 
Larix decidua, 255 
laricina, 404,574 
Lark, Horned, 71 
Larsen, K.W., 296 
Larus spp., 202 
argentatus, 203 
californicus, 203 
canus, 203 
delawarensis, 69 
glaucescens, 203 
philadelphia, 203 
thayeri, 203 
Larus spp., Post-Reproductive Pacific Salmon, Oncorhynchus 
spp., as a Major Nutrient Source for Large Aggrega- 
tions of Gulls, 202 
Lasallia, 78 
pensylvanica, 78 
Lasiurus cinereus, 132 
Lasiurus cinereus, Use of a Bridge for Day Roosting by the 
Hoary Bat, 132 
Lasius alienus, 40 
nearcticus, 40 
neoniger, 40 
niger, 40 
pallitarsis, 40 
umbratus, 40 
Lassioglossum sp., 249 
Lathyrus sphaericus, 248 
Lauff, R., Reviews by, 144,462 
Laurel, Sheep, 381 
Lavallée, B.F., 359 
Ledum decumbens, 210 
groenlandicum, 276,381 
Leersia oryzoides, 547 
Lefevre, K.L. Predation of a Bat by American Crows, Corvus 
brachyrhynchos, 443 
Lemming, Collared, 124 
Lenard: S., 132 
Lepisosteus osseus, 21 
Lepomis cyanellus, 21 
gibbosus, 21,360 
humilis, 21 
macrochirus, 21 
megalotis, 21 
Leproloma vouauxii, 78 
Leptarrhena pyrolifolia, 418 
Lepthyphantes spp., 263 
alpinus, 263 
bihamatus, 263 
calcaratus, 263 
complicatus, 263 
cracens, 263 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


duplicatus, 263 
intricatus, 263 
leprosus, 263 
nigriventris, 263 
subalpinus, 263 
tenuis, 263 
triramus, 263 
turbatrix, 263 
umbraticola, 263 
washingtoni, 263 
zebra, 263 

Leptothorax ambiguus, 39 
canadensis, 39 
ergatogyneus, 38 
longispinosus, 39 

Lepus americanus, 43,121,197,323,395,569 
arcticus, 323 
europaeus, 122 

Lepus americanus, on Insular Newfoundland During a Cycli- 
cal Peak, Analysis of the Parasites of a Mid-winter 
Population of the Snowshoe Hare, 323 

Lepus americanus, on Kent Island, New Brunswick, Tree 
Recruitment Limitation by Introduced Snowshoe 
Hares, 569 

Leslie, J.K. and C.A. Timmins. Environment and Distribution 
of Age 0 Fishes in River Canard, a Lowland Ontario 
River, 16 

Lestes disjunctus, 234 

Lethocerus sp., 292 

Leucorrhinia spp., 234 
intacta, 234 
proxima, 234 

Leymus mollis ssp. mollis, 417 

Libellula pulchella, 234 
quadrimaculata, 234 

Lichen, 413 

Lichter, J., 569 

Lily, Bullhead, 429 
Chocolate, 248 
White Fawn, 248 

Limosella aquatica, 277 

Linaria vulgaris, 14 

Linnaea borealis var. americana, 277 

Linyphia marginata, 264 
waldea, 264 

Liquorice-root, 287 

Lithophragma parviflorum, 248 

Little Nightmare, Eschscholtz’, 429 

Lizard, 95 

Lobelia inflata 394 

Locky, D.A., J.C. Davies and B.G. Warner. Effects of Wetland 
Creation on Breeding Season Bird Use in Boreal East- 
ern Ontario, 64 

Locoweed, Blackish, 432 

Logperch, 21 

Lolium spp., 548 

Lomatium utriculatum, 248 

Londry, K.L., PH. Badiou, and S.E. Grasby. Identification 
of a Marine Green Alga Percursaria percursa from 
Hypersaline Springs in the Middle of the North 
American Continent, 82 

Longspur, Lapland, 208 

Longspur, Calcarius lapponicus, Nests at Cape Churchill, 
Manitoba, Microhabitat Characteristics of Lapland, 
208 


2006 


Longue, P., 294 

Lonicera involucrata, 418 

Lontra canadensis, 459 

Lontra canadensis) Contribute to Fall Scent Marking?, Do 
Juvenile Nearctic River Otters, 459 

Loon, Common, 455 

Loon, Gavia immer, Nest Attendance Patterns Recorded by 
Remote Video Camera, Common, 455 

Looper, Hemlock, 324 

Loosestrife, Purple, 18 

Lophodytes cucullatus, 69 

Lophomma sylvaticum, 263 

Lotus, American, 17 

Lousewort, Lapland, 289 

Loxia curvirostra, 121 
leucoptera, 70 

Luetkea pectinata, 418 

Lumsden, H.G. Plumage and Internal Morphology of the 
“Prairie Grouse”, Tympanuchus cupido X phasianellus, 
of Manitoulin Island, Ontario, 515 

Lumsden, H.G. “Prairie Grouse”, Tympanuchus cupido x 
Phasianellus, Hybridization on Manitoulin Island, 
Ontario, 507 

Lupine, Nootka, 432 

Lupinus arcticus, 277 
nootkatensis, 418 

Lutra canadensis, 323 
lutra, 459 

Luxilus cornutus, 21,361 

Luzula piperi, 418 
spicata, 418 
wahlenbergii, 277 

Lycopodium sabinifolium ssp. sitchense, 418 
selago, 277 ' 
sitchense, 417 

Lycopus uniflorus, 277,388 

Lynx canadensis, 323 

Lynx, 323 

Lysimachia thyrsifolia, 389 

Lythrum salicaria, 18 


MaclInnes, K.I., 497 

Mack, C.M., 446 

MacPherson, A., Review by, 147 

Macrobiotus cf. areolatus, 587 
cf. hufelandi, 586 
cf. islandicus, 587 
cf. richtersi, 586 

Madsen, J.D., 546 

Madtom, Tadpole, 18 

Maianthemum canadense, 240 

Mallard, 379,541,546 

Mallards, Anas platyrhynchos, Use of Radio-Telemetry to 
Test for Investigator Effects on Nesting, 541 

Mallory, M.L., A.J. Fontaine, and H. Boyd. Breeding and 
Non-Breeding Range of Canada, Branta canadensis, 
and Cackling Geese, Branta hutchinsii, in the Eastern 
Canadian Arctic, 483 

Mallotus villosus, 216,355 

Mallow, Dwarf, 439 

Malva neglecta, 439 
rotundifolia, 438 

Manitoba, Microhabitat Characteristics of Lapland Longspur, 
Calcarius lapponicus, Nests at Cape Churchill, 208 


INDEX TO VOLUME 119 


633 


Mann, H. and E.M.V. Nambudiri. Charophytes of Insular 
Newfoundland II: Chara evoluta and Chara canes- 
cens, 26 

Maple, Manitoba, 220 
Red, 17,386 
Sugar, 38,332,378,386 

Mare’s-tail, 28 

Margariscus margarita, 221,360 

Marine Turtle Newsletter, 156,319,473,613 

Maro amplus, 263 
nearcticus, 263 

Marsh, J., 76 

Marsh-marigold, 285 
Floating, 285 

Marten, 124 
American, 43 
Newfoundland, 43 
Pine, 121,194,323 

Martens, Martes americana atrata, Seasonal Diets of New- 
foundland, 43 

Martes americana, 43,124,194,323 
americana atrata, 43 
martes, 121 
pennanti, 120,177,194 

Martes americana atrata, Seasonal Diets of Newfoundland 
Martens, 43 . 

Martin, P.A., S.R. de Solla, P.J. Ewins, and M.E. Barker. 
Productivity of Osprey, Pandion haliaetus, Nesting 
on Natural and Artificial Structures in the Kawartha 
Lakes, Ontario, 1991-2001, 58 

Mary, Blue-eyed, 248 

Maskinongé, 364 

Maso sundevallii, 263 

Matricaria ambigua, 277 

McCarthy, J., Reviews by, 153,308,316,601 

McGee, B.K., 584 

McGrath, M., 323 

McKague, C.I. and N. Cappuccino. Response of Pale Swal- 
low-wort, Vincetoxicum rossicum, following Above- 
ground Tissue Loss: Implications for the Timing of 
Mechanical Control, 525 

McMillan, B.R., 546 

McNay, M.E. and P.W. Mooney. Attempted Predation of a 
Child by a Gray Wolf, Canis lupus, near Icy Bay, 
Alaska, 197 

Meadow-foxtail, Water, 421 

Meadowlark, Eastern, 71 

Mecynargus paetulus, 263 

Medicago spp., 175,551 

Megachile sp., 250 
rotundata, 53 

Megaptera novaeangliae, 214 

Megascops asio, 578 

Meioneta sp. rurestris, 259 

Melampyrum lineare, 240 

Melandrium apetalum ssp. arcticum, 284 

Melanelia, 78 
disjuncta, 78 
sorediata, 78 
stygia, 78 

Melanerpes carolinus, 369 
erythrocephalus, 369 

Melanitta spp., 181 
fusca, 182 


634 


nigra, 182 
perspicillata, 182 
Melanitta spp., Distributions in Central Alaska Beaufort Sea 
Lagoons, 1999-2002, Long-tailed Duck, Clangula 
hyemalis, Eider, Somateria spp., and Scoter, 181 
Melospiza georgiana, 69 
lincolnii, 70 
melodia, 70,120 
melodius, 578 
Méné a nageoires rouges, 361 
laiton, 360 
Menyanthes trifoliata, 418 
Mephitis mephitis, 454,584 
Merchant, P.J., 580 
Merganser, Common, 69 
Hooded, 69 
Mergus merganser, 69 
Merlin, 71,379 
Merluccius productus, 112 
Merodon equestris, 249 
Meta menardi, 268 
ovalis, 268 
Metaphidippus flavipedes, 268 
montanus, 268 
Metasyrphus, 250 
Metepeira palustris, 257 
Metridium senile, 355 
Meunier noir, 360 
Micaria aenea, 258 
constricta, 259 
longispina, 259 
pulicaria, 259 
rossica, 259 
Mice, Peromyscus maniculatus, in Wire Mesh Live-Traps: 
A Cautionary Note, Mortality of Deer, 445 
Micrargus longitarsus, 263 
Microlinyphia mandibulata, 263 
Micropterus dolomieu, 21,295,353,363 
salmoides, 21,363 
Microstictia, 6 
Microtus spp., 122,395 
chrotorrhinus, 412 
chrotorrhinus chrotorrhinus, 412 
chrotorrhinus ravus, 412 
longicaudus, 445 
pennslyvanicus, 43,323,414,446 
xanthognathus, 446 
Microtus chrotorrhinus, in Labrador, A Range Extension for 
the Rock Vole, 412 
Milfoil, Eurasian, 16 
Milk-vetch, Alpine, 431 
Milkweed, 525 
Milnesium tardigradum, 586 
Miner’s-lettuce, 248 
Mink, 175 
Minnesota, Food Habits of Dabbling Ducks During Fall 
Migration in a Prairie Pothole System, Heron Lake, 
546 
Minnow, Bluntnose, 18 
Brassy, 219 
Eastern Silvery, 219 
Fathead, 21,221,351 
Mississippi Silvery, 219 
Plains, 219 
Pugnose, 18 
Silvery, 219 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Western Silvery, 219 
Minnow, Hybognathus placitus, in Canada, First Record of 
the Plains, 219 
Minnow, Pimephales promelas, in New Brunswick, The Fat- 
head, 351 
Minuartia biflora, 276 
dawsonensis, 418 
rossii, 276 
rubella, 277 
yukonensis, 277 
Minyriolus castaneus, 266 
Misumena vatia, 270 
Mniotilta varia, 70 
Moneses uniflora, 276 
Monotropa hypopithys, 240 
Montia linearis, 248 
Montia, Narrow-leaved, 248 
Mooney, P.W., 197 
Moose, 43,50,136,168,186,194,197,323,330,381,574 
Moose, Alces alces, and Beaver, Castor canadensis, in 
Algonquin Park, Ontario, 1860-2004, Reconstruct- 
ing Changes in Abundance of White-tailed Deer, 
Odocoileus virginianus, 330 
Moose, Alces alces, in Central Interior British Columbia, 
Winter Habitat Use by, 186 
Morone americana, 21,295 
chrysops, 21 
Mosgovoyia pectinata, 324 
Moss, 210,413,586 
Mountain-avens, White, 210 
Mountain-heather, Alaskan, 433 
Mountain-parsley, 278 
Mouse, 589,591 
Cotton, 446 
Deer, 194,323,414,445,451 
Grasshopper, 95 
House, 446,578 
Jumping, 578 
White-footed, 446,452,578 
Woodland, 578 
Mouse, Peromyscus maniculatus, in the Boreal Forest, Long- 
distance Movement of a Dispersing Deer, 451 
Mouse-ear, Little, 248 
Moxostoma sp., 21 
macrolepidotum, 21 
Mudminnow, Central, 19 
Mudwort, 288 
Aleutian, 435 
Muhlenbergia glomerata, 277 
Muldoon, F., 453 
Mulet a cornes, 361 
perlé, 360 
Multiceps sp., 324 
serialis, 324 
Murray, M.P., C.A. Pear, and R.B. Bury. Apparent Predation 
by Gray Jays, Perisoreus canadensis, on Long-toed 
Salamanders, Ambystoma macrodactylum, in the Ore- 
gon Cascade Range, 291 
Mus domesticus, 446 
musculus, 578 
Museau noir, 361 
Muskox, 168 
Muskrat, 570,591 
Mustela erminea, 175,323 
frenata, 175 
vison, 175 


2006 


Mustela frenata, Movements and Diggings in Alfalfa Fields 
Inhabited by Northern Pocket Gophers, Thomomys 
talpoides, Long-tailed Weasel, 175 

Myiarchus crinitus, 367 

Myosotis discolor, 248 

Myotis lucifugus, 132,444,589 

Myotis, Little Brown, 132 

Myotis lucifugus, in a Rodent Trap in the Boreal Forest, Mor- 
tality of Little Brown Bats, 589 

Myriophyllum sp., 16 
alterniflorum, 277 
verticillatum, 418 

Myrmecina americana, 39 

Myrmica detritinodis, 39 
emeryana, 39 
fracticornis, 39 
pinetorum, 39 
punctiventris, 39 
sculptilis, 39 

Mythoplastoides exiguus, 264 


Najas flexilis, 276,417 

Nambudiri, E.M.V., 26 

Neave, P., 453 

Nedra ramosula, 12 

Nelumbo lutea, 17 

Nematode, 324 

Némopanthe mucroné, 129 

Nemopanthus mucronatus, 129 

Neoantistea agilis, 259 
magna, 259 
riparia radula, 259 

Neogobius melanostomus, 21,582 

Neogobius melanostomus, in the St. Lawrence River at Corn- 
wall, Ontario, First Occurrence of the Round Goby, 
582 

Neon nellii, 268 

Neottiura bimaculata, 269 

Neriene clathrata, 264 
radiata, 264 

Nero, R.W. Short-tailed Shrew, Blarina brevicauda, Apparent- 
ly Killed by Red Squirrel, Tamiasciurus hudsonicus, 
456 

Nesticus pallida, 267 

Nevin, O.T. and B.K. Gilbert. Observations of Autumn 
Courtship and Breeding in Brown Bears, Ursus arc- 
tos, from Coastal British Columbia, 449 

New Brunswick, The Fathead Minnow, Pimephales prome- 
las, in, 351 

New Brunswick, Tree Recruitment Limitation by Introduced 
Snowshoe Hares, Lepus americanus, on Kent Island, 
569 

Newfoundland, An Annotated Checklist of the Spiders of, 
254 

Newfoundland During a Cyclical Peak, Analysis of the Par- 
asites of a Mid-winter Population of the Snowshoe 
Hare, Lepus americanus, on Insular, 323 

Newfoundland Martens, Martes americana atrata, Seasonal 
Diets of, 43 

Newfoundland II: Chara evoluta and Chara canescens, Charo- 
phytes of Insular, 26 

New York, Chronology of Range Expansion of the Coyote, 
Canis latrans, in, | 

Nicholls, K.H. Cyclopyxis acmodonta n. sp. and Arcella for- 
mosa n. sp.: Two New Species of Testate Rhizopods 


INDEX TO VOLUME 119 


635 


(Arcellinida, Protozoa) from Remnant Wetlands in 
Ontario, Canada, 403 

Nicholson, K.L., B.K. McGee, and W.B. Ballard. Swift Fox, 
Vulpes velox, Den Located Next to a Railroad Track 
in Northwestern Texas, 584 

Nitella, 26 
confervacea, 35 
flexilis, 35 
opaca, 35 

Noel, L.E., S.R. Johnson, and G.M. O’ Doherty. Long-tailed 
Duck, Clangula hyemalis, Eider, Somateria spp., and 
Scoter, Melanitta spp., Distributions in Central Alaska 
Beaufort Sea Lagoons, 1999-2002, 181 

Noisetier a long bec, 457 

North America (Coleoptera: Chrysomelidae, Chrysomelinae), 
New Distribution Records and Biogeography of Cal- 
ligrapha Species (Leaf Beetles), in, 88 

North American Banding, Long-eared Owls, Asio otus: A 
Review of, 395 

North American Continent, Identification of a Marine Green 
Alga Percursaria percursa from Hypersaline Springs 
in the Middle of the, 82 

Northwest Territories and Nunavut, Canada III, Additions 
and Range Extensions to the Vascular Plant Flora of 
the Continental, 276 

Northwest Territories from the Great Slave Lake Area, Addi- 
tions to the Flora of the Continental, 437 

Northwest Territories Links Two Allopatric Species of Alkali 
Grass, Puccinellia, Addition to the Flora of Canada? 
A Specimen from the Arctic Archipelago, 497 

Notemigonus crysoleucas, 21,295,361 

Notropis anogenus, 21 
atherinoides, 21 
heterolepis, 361 
hudsonius, 21 
stramineus, 21 
volucellus, 21 

Noturus gyrinus, 21 

Nova Scotia, Invasive Scots Pine, Pinus sylvestris, Replacing 
Corema, Corema conradii, Heathland in the Annapo- 
lis Valley, 237 

Nucifraga columbiana, 291 

Numenius americanus, 490 

Nunavut, Canada III, Additions and Range Extensions to the 
Vascular Plant Flora of the Continental Northwest Ter- 
ritories and, 276 

Nuphar variegatum, 388,418 

Nutcracker, Clark’s, 291 

Nuthatch, Red-breasted, 71,367 
White-breasted, 367 

Nymph, Wavy Water, 279,420 

Nymphaea tetragona ssp. leibergii, 277 


O’ Doherty, G.M., 181 

O’ Donovan, K.S., 445,451 

O'Neill, J., Reviews by, 463,464 

Oak, 343,368 
Bur, 348 
Garry, 245,344 
Red, 38,348,386 
Scrub, 9 

Oak Forests, Woodpecker Nest Tree Characteristics in Upper 
Midwestern, 367 

Oak, Quercus garryana, Ecosystem of British Columbia, 
Insect Visitation to Wildflowers in the Endangered 
Garry, 245 


636 


Oatgrass, Poverty, 279 

Oats, 421 

Obbard, M., 48 

Obeliscoides cuniculi, 324 

Oceanodroma leucorhoa, 570 

Odocoileus heminous, 296,574,580 
virginianus, 101,330,381,573,580,591 

Odocoilus hemionus, and Other Mammals, Passage Through 
a Small Drainage Culvert by Mule Deer, 296 

Odocoileus virginianus, Moose, Alces alces, and Beaver, 
Castor canadensis, in Algonquin Park, Ontario, 
1860-2004, Reconstructing Changes in Abundance 
of White-tailed Deer, 330 

Oedothorax montiferus, 266 
trilobatus, 264 

Olson, R.W., J.K. Schmutz, and U.T. Hammer. Occurrence, 
Composition and Formation of Ruppia, Widgeon 
Grass, balls in Saskatchewan Lakes, 114 

Olson, Z.H., S.S. Stevens, and T.L. Serfass. Do Juvenile 
Nearctic River Otters (Lontra canadensis) Contribute 
to Fall Scent Marking? 459 

Ommatokoita elongata, 355 

Oncorhynchus spp., 202 
gorbuscha, 202 
keta, 202 
kisutch, 197 
tshawytscha, 591 

Oncorhynchus spp., as a Major Nutrient Source for Large 
Aggregations of Gulls, Larus spp., Post-Reproductive 
Pacific Salmon, 202 

Ondatra zibethicus, 570,591 

Ontario: An Addition to the Flora of Canada, Papillate Water- 
meal, Wolffia brasiliensis, in Eastern, 137 

Ontario, Canada, Cyclopyxis acmodonta n. sp. and Arcella 
formosa n. sp.: Two New Species of Testate Rhi- 
zopods (Arcellinida, Protozoa) from Remnant Wet- 
lands in, 403 

Ontario, Effects of Wetland Creation on Breeding Season 
Bird Use in Boreal Eastern, 64 

Ontario, First Occurrence of the Round Goby, Neogobius 
melanostomus, in the St. Lawrence River at Cornwall, 
582 

Ontario, Identification and Status of the Introduced Black 
Pine, Pinus nigra, and Mugo Pine, Pinus mugo, in, 
224 

Ontario, Movements of Two Rabid Raccoons, Procyon lotor, 
in Eastern, 453 

Ontario Natural Heritage Information Centre Science and 
Information Newsletter, 319 

Ontario, 1860-2004, Reconstructing Changes in Abundance 
of White-tailed Deer, Odocoileus virginianus, Moose, 
Alces alces, and Beaver, Castor canadensis, in Algon- 
quin Park, 330 

Ontario, Plumage and Internal Morphology of the “Prairie 
Grouse”, Tympanuchus cupido X phasianellus, of 
Manitoulin Island, 515 

Ontario, “Prairie Grouse”, Tympanuchus cupido X phasianel- 
lus, Hybridization on Manitoulin Island, 507 

Ontario, Productivity of Osprey, Pandion haliaetus, Nesting 
on Natural and Artificial Structures in the Kawartha 
Lakes, Ontario, 1991-2001, 58 

Ontario River, Environment and Distribution of Age 0 Fishes 
in River Canard, a Lowland, 16 

Ontario Watershed, Possible Tool Use by Beavers, Castor 
canadensis, in a Northern, 441 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Ontario Woodlands, Detectability of Non-passerines Using 
“Pishing” in Eastern, 377 
Onychomys, 95 
Oporornis agilis, 70 
philadelphia, 70 
Opsopoeodus emiliae, 21 
Opuntia polyacantha, 220 
Orchid, Bracted Green, 428 
Orconectes propinquus, 17 
Oreamnos americanus, 197,574 
Oregon Cascade Range, Apparent Predation by Gray Jays, 
Perisoreus canadensis, on Long-toed Salamanders, 
Ambystoma macrodactylum, in the, 291 
Oreonetides spp., 264 
flavescens, 264 
rectangulatus, 264 
rotundus, 264 
vaginatus, 264 
Oreophantes recurvatus, 264 
Oriole, Baltimore, 70 
Orobanche fasciculata, 418 
Orodrassus canadensis, 259 
vastus, 259 
Osmia sp., 246 
lignaria, 246 
texana, 246 
Osprey, 58,379 
Osprey, Pandion haliaetus, Nesting on Natural and Artificial 
Structures in the Kawartha Lakes, Ontario, 1991- 
2001, Productivity of, 58 
Ostrya, 96 
Ottawa Field-Naturalists’ Club Awards for 2004, The, 614 
Ottawa Field-Naturalists’ Club 11 January 2005, Minutes of 
the 126'" Annual Business Meeting of The, 474 
Otter, European, 459 
Nearctic River, 459 
River, 323 
Otters (Lontra canadensis) Contribute to Fall Scent Mark- 
ing?, Do Juvenile Nearctic River, 459 
Ovenbird, 71 
Ovibos moschatus, 168 
Ovis canadensis, 574 
Owl, Barred, 367,379 
Boreal, 367,578 
Burrowing, 490 
Eastern Screech, 578 
Great Gray, 69 
Great Horned, 62,71,395,578 
Long-eared, 395 
Northern Saw-whet, 367,578 
Short-eared, 70,400 
Snowy, 211 
Owls, Aegolius acadicus, Caching Behavior by Wintering 
Northern Saw-Whet, 578 
Owls, Asio otus: A Review of North American Banding Long- 
eared, 395 
Oxyptila americana, 270 
Oxytrope, Arctic, 432 
Jordal’s, 432 
Murray’s, 432 
Oxytropis arctica var. arctica, 417 
arctica var. murrayi, 417 
arctobia, 277 
campestris ssp. jordalii, 418 
deflexa ssp. foliosa, 277 


2006 


maydelliana, 277 

nigrescens ssp. nigrescens, 418 

sordida ssp. murrayi, 432 
Ozyptila bryantae, 270 

distans, 270 

gertschi, 270 

sincera canadensis, 270 


Pachygnatha brevis, 268 
Pagophilus groenlandicus, 382 
Paintbrush, Scarlet, 433 
Small-flowered, 434 
Unalaska, 434 
Pandion haliaetus, 58,379 
Pandion haliaetus, Nesting on Natural and Artificial Struc- 
tures in the Kawartha Lakes, Ontario, 1991-2001, Pro- 
ductivity of Osprey, 58 
Panicum capillare var. occidentale, 439 
dichotomiflorum, 548 
miliaceum, 438 
Papaver radicatum, 277 
radicatum ssp. kluanensis, 418 
Papilio sp., 249 
Paquet, PC... 192 
Parachnowitsch, A.L. and E. Elle. Insect Visitation to Wild- 
flowers in the Endangered Garry Oak, Quercus gar- 
ryana, Ecosystem of British Columbia, 245 
Paralochloa, 497 
Pardosa albomaculata, 267 
concinna, 267 
furcifera, 267 
fuscula, 267 
groenlandica, 267 
hyperborea, 267 
lapidicina, 267 
mackenziana, 267 
moesta, 267 
saltuaria, 267 
xerampelina, 267 
Parelaphostrongylus tenuis, 334 
Parmelia, 78 
saxatilis, 78 
sulcata, 78 
Parnassia fimbriata, 418 
Parrya arctica, 277,506 
Partridge, Grey, 122 
Partridge-foot, 431 
Partridgeberry, 412 
Parula americana, 69 
Parus atricapillus, 367 
Passer domesticus, 71 
Passerculus sandwichensis, 70,570 
Pear, C.A., 291 
Peavine, Grass, 248 
Pedicularis arctica, 289 
capitata, 277,506 
langsdorfii ssp. arctica, 277,506 
lapponica, 277 
macrodonta, 277 
parviflora var. macrodonta, 289 
Pelecopsis mengei, 264 
Pelegrina flavipes, 268 
montana, 268 
Perca flavescens, 21,295 
Perch, White, 18,295 


INDEX TO VOLUME 119 


637 


Yellow, 20,295 
Percina caprodes, 21 
maculata, 21 
Percursaria percursa, 82 
Percursaria percursa from Hypersaline Springs in the Middle 
of the North American Continent, Identification of a 
Marine Green Alga, 82 
Perdix perdix, 122 
Perisoreus canadensis, 70,101,291 
Perisoreus canadensis: Adaptations to Boreal Winters, 
Weight-carrying Ability and Caching Behavior of 
Gray Jays, 101 
Perisoreus canadensis, on Long-toed Salamanders, Amby- 
stoma macrodactylum, in the Oregon Cascade Range, 
Apparent Predation by Gray Jays, 291 
Peromyscus sp., 446,452,578,589,59 1 
gossypinus, 446 
leucopus, 446,452,578 
maniculatus, 194,323,414,445,451 
Peromyscus maniculatus, in the Boreal Forest, Long-distance 
Movement of a Dispersing Deer Mouse, 451 
Peromyscus maniculatus, in Wire Mesh Live-Traps: A Cau- 
tionary Note, Mortality of Deer Mice, 445 
Peterson, T.S., A. Uesugi, and J. Lichter. Tree Recruitment 
Limitation by Introduced Snowshoe Hares, Lepus 
americanus, on Kent Island, New Brunswick, 569 
Petrel, Leach’s Storm, 570 
Petrochilidon pyrrhonota, 70 
Pewee, Eastern Wood, 71 
Phaeophyscia hispidula, 78 
sciastra, 78 
Phalacrocorax auritus, 69 
Phalaris arundinacea, 418 
Phenacomys intermedius, 414 
Pheucticus ludovicianus, 71 
Phidippus borealis, 268 
princeps, 268 
purpuratus, 268 
whitmanti, 268 
Philanthus albopilosus, 9 
politus, 10 
Phillips, RR., 381 
Philodromus alascensis, 268 
imbecillus, 268 
mysticus, 268 
pernix, 268 
placidus, 268 
rufus, 268 
rufus quartus, 268 
Phippsia algida, 418 
Phlox gracilis, 248 
Phlox, Slender, 248 
Phoca hispida, 357 
vitulina, 355 
Pholcus phalangioides, 268 
Phoxinus eos, 221,360 
neogaeus, 360 
Phragmites australis, 17,277 
communis var. berlandieri, 280 
Phyciodes mylitta, 249 
Phyllodoce xX intermedia, 418 
Physcia caesia, 78 
dubia, 78 
phaea, 78 
Physconia muscigena, 78 


638 


Physidae, 547 
Physocarpus, 96 
opulifolius, 96 
Picea sp., 135,208,457 
engelmanii X glauca, 186 
glauca, 77,186,255,381,404,45 1,458,574 
mariana, 186,255,333,381,412,457,574 
rubens, 570 
sitchensis, 160,197 
Pickavance, J.R. and C.D. Dondale. An Annotated Checklist 
of the Spiders of Newfoundland, 254 
Picoides arcticus, 367 
dorsalis, 367 
pubescens, 71,369,379 
villosus, 71,369,379 
Pieris rapae, 249 
Pike, Northern, 21 
Pimephales notatus, 21,361 
promelas, 21,221,351,361 
Pimephales promelas, in New Brunswick, The Fathead Min- 
now, 351 
Pin gris, 457 
rouge, 457 
Pine, Black, 224 
Eastern White, 386 
Jack, 332 
Japanese Black, 224 
Japanese Red, 224 
Lodgepole, 186,419,451,574 
Mugo, 224 
Pitch, 9 
Ponderosa, 132 
Red, 338,237,332 
Scotch, 8,231 
Scots; 231,237 
Shore, 344 
White, 9,38,238,332,369,378 
Pine, Pinus mugo, in Ontario, Identification and Status of the 
Introduced Black Pine, Pinus nigra, and Mugo, 224 
Pine, Pinus nigra, and Mugo Pine, Pinus mugo, in Ontario, 
Identification and Status of the Introduced Black, 
224 
Pine, Pinus sylvestris, Replacing Corema, Corema conradii, 
Heathland in the Annapolis Valley, Nova Scotia, Inva- 
sive Scots, 237 
Pinicola enucleator, 121 
Pintail, Northern, 69,548 
Pinus spp., 548 
banksiana, 231,332,457 
cembroides var. edulis, 231 
contorta, 186,344,451,574 
contorta ssp. latifolia, 418 
densiflora, 224 
edulis, 231 
montana, 229 
mugo, 224 
mugo ssp. mugo, 229 
mugo ssp. rotundata, 229 
mugo ssp. uncinata, 229 
nigra, 224 
nigra var. nigra, 226 
ponderosa, 132 
resinosa, 8,38,231,237,332,457 
rigida, 9 
strobus, 9,38,238,332,369,378,386 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


sylvestris, 8,231,237 
thunbergii, 224 
Pinus mugo, in Ontario, Identification and Status of the Intro- 
duced Black Pine, Pinus nigra, and Mugo Pine, 224 
Pinus nigra, and Mugo Pine, Pinus mugo, in Ontario, Identi- 
fication and Status of the Introduced Black Pine, 224 
Pinus sylvestris, Replacing Corema, Corema conradii, Heath- 
land in the Annapolis Valley, Nova Scotia, Invasive 
Scots Pine, 237 
Pipit, Sprague’s, 490 
Pirata bryantae, 267 
canadensis, 267 
cantralli, 267 
insularis, 267 
minutus, 267 
montanus, 267 
piraticus, 267 
Pityohyphantes costatus, 264 
limitaneus, 264 
subarcticus, 264 
Placidium squamulosum, 78 
Planorbidae, 547 
Plantago eriopoda, 418 
major, 394 
Plantain, Saline, 434 
Platanthera clavellata, 388 
Platycrista cheleusis, 587 
Plecoptera, 547 
Plectrophanax nivalis, 120 
Plenoculus davisi, 10 
Plethodon cinereus, 129 
Pletscher, D.H., 446 
Pleurozium schreberi, 586 
Plover, Mountain, 532 
Plovers, Charadrius montanus, Differential Parental Care by 
Adult Mountain, 532 
Poa alpigena, 280 
alpina, 277 
glauca, 277 
pratensis, 245 
pratensis ssp. alpigena, 277 
Pocadicnemis americana, 264 
Podagrostis thurberiana, 420 
Podiceps auritus, 69 
Podilymbus podiceps, 69 
Poecile spp., 121 
atricapillus, 71 
cinctus, 120 
hudsonica, 70,367 
major, 120 
montanus, 120 
palustris, 120 
Poeciloneta sp., 264 
furcata, 264 
Pogonia ophioglossoides, 388 
Polemonium pulcherrimum, 418 
Polites themistocles, 12 
Polygonum spp., 547 
achoreum, 418 
convolvulus, 418 
lapathifolium, 418,547 
pennsylvanicum, 548 
pensylvanicum ssp. oneillii, 428 
persicaria, 394,418,548 
viviparum, 277 


2006 


Polytrichum, 78 
lonchitis, 418 
Pomoxis annularis, 21 
nigromaculatus, 21 
Pondhawk, Eastern, 234 
Pondweed, Blunt-leaved, 279 
Clasping-leaved, 28 
Closed-leaved, 278 
Curlyleaf, 546 
Fine-leaved, 278 
Horned, 28 
Sago, 28 
White-stemmed, 420 
Ponera pennsylvanica, 39 
Poplar, 41 
Balsam, 77 
Populus sp., 41,367,401,457 
balsamifera, 77 
deltoides, 17,132 
tremuloides, 65,135,175,186,220,381,442,45 1,574 
Porcupine, 125,159,177,453 
Porcupines, Erethizon dorsatum, in Second-Growth Conifer 
Forests, The Influence of Thermal Protection on Win- 
ter Den Selection by, 159 
Porrhomma sp., 264 
gertschi, 264 
terrestre, 264 
Portulaca oleracea, 438 
Porzana carolina, 69 
Posidonia, 114 
Potamogeton sp., 16,546 
crispus, 546 
dichotomiflorum, 547 
epihydrus, 394 
filiformis, 277 
foliosus, 277 
foliosus var. macellus, 278 
natans, 548 
obtusifolius, 276 
pectinatus, 28 
perfoliatus, 28 
praelongus, 418 
pusillus, 547 
Potentilla anserina, 560 
biflora, 277 
bipinnatifida, 418 
nivea ssp. nivea, 277 
norvegica, 394 
palustris, 277 
recta, 394 
potentille ansérine, 560 
Pouliot, D., 105 
Pouliot, D. et J.-F. Desroches. Découverte de la Salamandre 
a quatre orteils, Hemidactylium scutatum, 4 Québec, 
Québec: limite nord-est de l’espéce sur la rive nord 
du fleuve Saint-Laurent, 129 
Powell, T., 451 
Prairie-dog, Black-tailed, 532 
Price, M.H.H., C.T. Darimont, N.N. Winchester, and P.C. 
Paquet. Facts from Faeces: Prey Remains in Wolf, 
Canis lupus, Faeces Revise Occurrence Records for 
Mammals of British Columbia’s Coastal Archipelago, 
192 
Primrose, Siberian, 433 
Primula nutans, 418 


INDEX TO VOLUME 119 


639 


sibirica, 433 
Proctor, H.C., 586 
Procyon lotor, 62,298,453 
Procyon lotor, in Eastern Ontario, Movements of Two Rabid 
Raccoons, 453 
Proietto, R.L., 139 
Proulx, G. Body Weights of Adult and Juvenile Northern 
Pocket Gophers, Thomomys talpoides, in Central Al- 
berta Alfalfa Fields, 551 
Proulx, G. Long-tailed Weasel, Mustela frenata, Movements 
and Diggings in Alfalfa Fields Inhabited by North- 
ern Pocket Gophers, Thomomys talpoides, 175 
Proulx, G. and R.M. Kariz. Winter Habitat Use by Moose, 
Alces alces, in Central Interior British Columbia, 186 
Pruche du Canada, 129 
Pruitt, W.O., Jr., Why and How to Study a Snowcover, 118 
Prunus spp., 344 
Psathyrostachys juncea, 418 
Pseudacris triseriata, 291 
Pseudocolpodium, 498 
Pseudotsuga menziesii, 344 
Ptarmigan, Willow, 44 
Pteridium aquilinum, 240 
Pterostichus melanarius, 271 
Puccinellia, 497 
agrostidea, 498 
angustata, 497 
arctica, 497 
beringensis, 498 
byrrangensis, 498 
colpodioides, 498 
Jenisseiensis, 498 
phryganodes, 497 
poacea, 498 
vahliana, 498 
vahliana X, 498 
wrighti var. colpodioides, 498 
wrighti var. flava, 498 
wrighti var. wrightii, 498 
wrightii, 498 
Puccinellia, Addition to the Flora of Canada? A Specimen 
from the Arctic Archipelago, Northwest Territories 
Links Two Allopatric Species of Alkali Grass, 497 
Puma concolor, 580 
Puma concolor, in the Yukon, First Confirmation of Cougar, 
580 
Pumpkinseed, 21 
Pusa hispida, 382 
Pusillia mandibulata, 263 
Pussytoes, Showy, 434 
Pyrola secunda, 277 
Pyrola, One-flowered, 288 
Pyrrhula pyrrhula, 120 


Quebec, Consequences of Beaver, Castor canadensis, Flood- 
ing on a Small Shore Fen in Southwestern, 385 

Québec, et impact sur l’ichtyofaune, Charactéristiques d’ une 
population introduit du Grand brochet, Esox lucius, 
dans le lac Ramsay, Pare de la Gatineau, 359 

Québec, Extension de l’aire de distribution de la Couleuvre 
a collier, Diadophis punctatus edwardsii, dans lest 
du, 457 

Québec: limite nord-est de l’espéce sur la rive nord du fleuve 
Saint-Laurent, Découverte de la Salamandre a qua- 
tre orteils, Hemidactylium scutatum, a Québec, 129 


640 


Québec, Premiéres mentions et répartition de la Salamandre 
sombre du Nord, Desmognathus fuscus, sur la rive 
nord du fleuve Saint-Laurent, au, 105 

Québec, Québec: limite nord-est de l’espéce sur la rive nord 
du fleuve Saint-Laurent, Découverte de la Salamandre 
a quatre orteils, Hemidactylium scutatum, a, 129 

Québec, The Effect of Human Activity on Ant Species 
(Hymenoptera: Formicidae) Richness at the Mont 
St. Hilaire Biosphere Reserve, 38 

Quercus spp., 9,343 
alba, 368 
bicolor, 368 
garryana, 245,344 
macrocarpa, 348 
rubra, 38,129,240,348,368,386 

Quercus garryana, Ecosystem of British Columbia, Insect 
Visitation to Wildflowers in the Endangered Garry 
Oak, 245 

Quillback, 18 

Quinn, N.W.S. Reconstructing Changes in Abundance of 
White-tailed Deer, Odocoileus virginianus, Moose, 
Alces alces, and Beaver, Castor canadensis, in Algon- 
quin Park, Ontario, 1860-2004, 330 

Quiscalus quiscula, 70 


Raccoon, 62,298,453 
Raccoons, Procyon lotor, in Eastern Ontario, Movements of 
Two Rabid, 453 
Rail, Virginia, 69 
Yellow, 69 
Rallus limicola, 69 
Ramalina intermedia, 78 
Ramazzottius sp., 586 
Rana luteiventris, 291 
palustris, 578 
pipiens, 389 
Rangifer tarandus, 43,119,168,201,355,381 
tarandus caribou, 124,136,573 
tarandus fennicus, 124 
tarandus granti, 124 
tarandus groenlandicus, 124 
tarandus pearyi, 120 
tarandus tarandus, 124 
Ranunculus aquatilis var. eradicatus, 277 
occidentalis, 248 
repens, 438 
sabinei, 277 
xspitzbergensis, 277 
Raspberry, 369 
Common, 44 
Red, 570 
Wild Red, 287 
Rattlesnake, 95 
Rauschia triangularis, 324 
Raven, Common, 70,291,444 
Reading, K.L., 276 
Reddoch, A.H., 385 
Reddoch, J.M. and A.H. Reddoch. Consequences of Beaver, 
Castor canadensis, Flooding on a Small Shore Fen 
in Southwestern Quebec, 385 
Redhorse sp., 21] 
Shorthead, 21 
Redpoll, 120 
Common, 120 
Redstart, American, 71 
Reed, Common, 17,280 


THE CANADIAN FIELD-NATURALIST 


Vol 149 


Reedgrass, Bluejoint, 77 
Reeves, H.M., Review by, 307 
Regulus calendula, 70 
satrapa, 579 
Reimchen, T.E., 202 
Rheum rhabarbarum, 438 
Rhinichthys cataractae, 221 
Rhizocarpon disporum, 78 
Rhizoplaca chrysoleuca, 78 
Rhododendron groenlandicum, 412 
lapponicum, 210 
Rhynchospora alba, 277 
Rhytidium rugosum, 80 
Ribes spp., 369 
Robertus banksi, 269 
borealis, 269 
fuscus, 269 
riparius, 269 
Robin, American, 70 
Rogers, L.L. Weight-carrying Ability and Caching Behavior 
of Gray Jays, Perisoreus canadensis: Adaptations to 
Boreal Winters, 101 
Rohner, C., 573 
Rosa spp., 176 
blanda, 277 
woodsii, 418 
Rosatte, R., M. Allan, R. Warren, P. Neave, T. Babin, L. 
Buchanan, D. Donovan, K. Sobey, C. Davies, F. 
Muldoon, and A. Wandeler. Movements of Two Rabid 
Raccoons, Procyon lotor, in Eastern Ontario, 453 
Rose, 176 
Western, 431 
Rosebay, Lapland, 210 
Rosemary, Bog, 210 
Roussel, B., 457 
Roy, J.F., Review by, 605 
Rubus spp., 239,369 
arcticus ssp. acaulis, 277 
chamaemorus, 210,412 
idaeus, 44,277,570 
strigosus, 287 
vermontanus, 240 
Rugathodes aurantius, 269 
sexpunctatus, 269 
Rumex acetosella, 240 
crispus, 438 
maritimus ssp. fueginus, 418 
Ruppia, 114 
cirrhosa, 115 
maritima, 28,115 
occidentalis, 115 
Ruppia, Widgeon Grass, balls in Saskatchewan Lakes, Occur- 
rence, Composition and Formation of 114 
Rush, Bog, 282 
Swaying, 422 
Thread, 427 
Toad, 427 
Rye, 422 
Canadian Wild, 279 
Russian Wild, 422 


Sagebrush, 220,532 
Sagina caespitosa, 277 
nivalis, 418 
Sagittaria, 390 
latifolia, 394,548 


2006 


Salad, Corn, 248 
Salamander, Long-toed, 291 
Northwestern, 292 
Salamanders, Ambystoma macrodactylum, in the Oregon 
Cascade Range, Apparent Predation by Gray Jays, 
Perisoreus canadensis, on Long-toed, 291 
Salamandre a deux lignes, 106 
a quatre orteils, 129 
cendrées , 129 
pourpre, 106 
sombre du Nord, 105 
Salamandre a quatre orteils, Hemidactylium scutatum, a 
Québec, Québec: limite nord-est de l’espéce sur la 
rive nord du fleuve Saint-Laurent, Découverte de la, 
129 
Salamandre sombre du Nord, Desmognathus fuscus, sur la rive 
nord du fleuve Saint-Laurent, au Québec, Premiéres 
mentions et répartition de la, 105 
Salix sp., 65,77,96, 132,135,175, 186,208,220,44 1 506,574,591 
alaxensis, 277 
arctica, 277 
arctica X glauca, 276 
arctophila, 210,277 
bebbiana, 96 
brachycarpa ssp. niphoclada, 428 
fuscescens, 277 
glauca ssp. callicarpaea, 276 
glauca var. acutifolia, 277 
lanata ssp. richardsonii, 283 
myrtillifolia, 276 
niphoclada, 418 
planifolia, 277 
reticulata, 277 
richardsonii, 277 
rotundifolia, 276 
rotundifolia ssp. rotundifolia, 283 
sphenophylla, 277 
Salmon, 591 
Chinook, 591 
Chum, 202 
Coho, 197 
Pacific, 202 
Pink, 202 
Salmon Carcasses: Opportunistic Use of a Seasonally Super- 
abundant Food Source, Beavers, Castor canadensis, 
Feeding on, 591 
Salmon, Oncorhynchus spp., as a Major Nutrient Source for 
Large Aggregations of Gulls, Larus spp., Post-Repro- 
ductive Pacific, 202 
Salmonid, 353 
Salticus scenicus, 268 
Salvelinus fontinalis, 351 
Sanderson, E.W., | 
Sandpiper, Least, 379 
Spotted, 69,379 
Upland, 69 
Sandwort, Bog, 429 
Sanicle, Pacific, 248 
Sanicula crassicaulis, 248 
Sapin baumier, 129,457 
Sapsucker, Yellow-bellied, 71,367,379 
Sarcobatus vermiculatus, 220 
Sarsparilla, Wild, 44 
Saskatchewan Lakes, Occurrence, Composition and Forma- 
tion of Ruppia, Widgeon Grass, balls in, 114 
Saskatoon, 78 


INDEX TO VOLUME 119 


641 


Satilatlas gertschi, 264 
marxii, 264 
Savannah Sparrow, 70 
Saving the Wild: RENEW 2005, 613 
Saxifraga nelsoniana ssp. carlottae, 417 
nelsoniana ssp. pacifica, 418 
nelsoniana ssp. porsildiana, 277 
nivalis, 277 
oppositifolia, 506 
punctata ssp. carlottae, 431 
punctata ssp. pacifica, 431 
punctata ssp. porsildiana, 286 
rivularis, 277 
rufopilosa, 418 
tricuspidata, 78,506 


Saxifrage, Alpine, 286 
Leather-leaved, 430 
Prickly, 78 
Schmiegelow, F.K.A., 573 
Schmutz, J.K., 114 
Schoenoplectus acutus, 418 
acutus var. acutus, 427 
pungens, 439 
tabernaemontani, 418 
Sciastes dubius, 262 
truncatus, 264 
Scirites pectinatus, 264 
Scironis tarsalis, 265 
Scirpus sp., 16 
acutus, 418,548 
americanus, 439 
caespitosus ssp. austriacus, 277 
cyperinus, 137,394 
fluviatilis, 548 
hudsonianus, 418 
rollandii, 277 
rufus, 417 
validus, 17,418,548 
Sciurus carolinensis, 343,456 
vulgaris, 121,343 
Sciurus carolinensis, in British Columbia, The Distribution 
and Habitat Selection of Introduced Eastern Grey 
Squirrels, 343 
Scolochloa festucacea, 77 
Scolopax minor, 69 
Scoter, 181 
Black, 182 
Surf, 182 
White-winged, 182 
Scoter, Melanitta spp., Distributions in Central Alaska Beau- 
fort Sea Lagoons, 1999-2002, Long-tailed Duck, Clan- 
gula hyemalis, Eider, Somateria spp., and, 181 
Scotinotylus pallidus, 265 
sacer, 265 
Sculpin, 582 
Scutellaria galericulata, 394 
Scyletria inflata, 265 
Seal, Grey, 355 
Harbour, 355 
Harp, 382 
Hooded, 382 
Ringed, 357,382 
Seburn, D., Review by, 145 
Secale cereale, 418 
Sedge, 16,186,211,591 


642 


Awned, 77 
Bebb’s, 423 
Blackish, 426 
Bristle-leaved, 424 
Bristly, 137 
Brownish, 423 
Buxbaum’s, 423 
Dunhead, 426 
Enander’s, 425 
Few-seeded Fen, 425 
Grey, 424 
Hay, 426 
Kellogg’s, 425 
Pale, 425 
Parry’s, 426 
Sartwell’s, 426 
Slender, 424 
Small-winged, 425 
Spikehead, 426 
Weak, 425 
Seiurus aurocapillus, 71 
noveboracensis, 69 
Semljicola obtusus, 265 
Semotilus atromaculatus, 295,351,361 
Senecio congestus, 277 
Senfass,, T.L...459 
Sericocarpus rigidus, 245 
Setaria spp., 548 
glauca, 438 
Setophaga ruticilla, 71 
Shad, Gizzard, 18 
Shark, Bluntnose Sixgill, 537 
Greenland, 355 
Sharks, Hexanchus griseus, at Flora Islets, British Columbia, 
Body Size Distribution and Frequency of Anthro- 
pogenic Injuries of Bluntnose Sixgill, 537 
Sharks, Somniosus microcephalus, Vision and its Relation- 
ship to Novel Behaviour in St. Lawrence River Green- 
land, 355 
Sheep, 200 
Bighorn, 574 
Shepherdia azgenteas, 220 
canadensis, 276 
Shik, J.Z., A. Francoeur, and C.M. Buddle. The Effect of 
Human Activity on Ant Species (Hymenoptera: For- 
micidae) Richness at the Mont St. Hilaire Biosphere 
Reserve, Québec, 38 
Shiner, Common, 21 
Emerald, 20 
Golden, 21,295 
Mimic, 21 
Pugnose, 18 
Sand, 21 
Spotfin, 18 
Spottail, 21 
Shootingstar, Broad-leaved, 248 
Shoveler, Northern, 69,548 
Shrew, 122,578 
Masked, 43,324,579 
Pygmy, 414 
Short-tailed, 456,578 
Shrew, Blarina brevicauda, Apparently Killed by Red Squir- 
rel, Tamiasciurus hudsonicus, Short-tailed, 456 
Shrike, 102,578 
Loggerhead, 490 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Silene acaulis ssp. subacaulescens, 418 
cserei, 439 
walensis, 277 
Silverside, Brook, 21 
Singa variabilis, 257 
Sisicottus montanus, 265 
quoylei, 265 
Sisicus apertus, 265 
penifusifer, 265 
Sisis rotundus, 265 
Siskin, Pine, 71 
Sitta canadensis, 71,367 
carolinensis, 367 
Sitticus cutleri, 268 
finschii, 268 
floricola palustris, 268 
palustris, 268 
ranieri, 268 
striatus, 268 
Skunk, 454 
Striped, 584 
Slough, B.G., 589 
Smilacina trifolia, 44 
Smith, D.G., 578 
Snake, Garter, 292 
Snap-dragon, Dwarf, 438 
Snipe, Wilson’s, 69 
Snowberry, Creeping, 44 
Soapberry, 287 
Sobey, K., 453 
Solenopsis molesta, 39 
Solidago sp., 388 
altissima, 525 
bicolor, 240 
canadensis, 390 
graminifolia var. major, 277 
Juncea, 240 
uliginosa, 388 
Solomon’s Seal, Three-Leaved False, 44 
Somateria spp., 181 
mollissima v-nigrum, 182 
spectabilis, 182 
Somateria spp., and Scoter, Melanitta spp., Distributions in 
Central Alaska Beaufort Sea Lagoons, 1999-2002, 
Long-tailed Duck, Clangula hyemalis, Eider, 181 
Somniosus microcephalus, 355 
Somniosus microcephalus, Vision and its Relationship to 
Novel Behaviour in St. Lawrence River Greenland 
Sharks, 355 
Sonchus arvensis ssp. uliginosus, 439 
arvensis var. glabrescens, 439 
uliginosus, 438 
Sora, 69 
Sorbus americanus, 570 
sitchensis, 418 
Sorex sp., 122,578 
cinereus, 43,324,579 
hoyi, 414 
Souessa spinifera, 265 
Sougambus bostoniensis, 265 
Soulgas corticarius, 265 
Soybean, 547 
Sparganium sp., 16,390 
americanum, 137,389 
hyperboreum, 277 


2006 


Sparrow, 578 
Chipping, 71 
House, 71 
LeConte’s, 69 
Lincoln’s, 70 
Savannah, 570 
Song, 70,120,578 
Swamp, 69 
White-crowned, 71 
White-throated, 70 
Spartina gracilis, 277 
Spermophilus richardsonii, 442 
Sphagnum sp., 129,388,404 
magellanicum, 388 
teres, 388 
Sphaigne, 129 
Sphixapata vigilans, 13 
Sphyrapicus varius, 71,367,379 
Spider, 95,254 
Spiders of Newfoundland, An Annotated Checklist of the, 
254 
Spike-rush, Bald, 439 
Spiraea betulifolia, 438 
Spirembolus oreinoides, 265 
Spizella passerina, 71 
Spreadwing, Northern, 234 
Spruce, 135,186,208 
Black, 186,255,333,381,412,574 
Red, 570 
Sitka, 160,197 
White, 77,186,255,381,404,45 1,570,574 
Squirrel, Douglas, 344 
Eastern Grey, 343 
European Red, 343 
Grey, 456 
North American Red, 121,343 
Northern Flying, 344,591 
Red, 43,121,323,456,591 
Richardson’s Ground, 442 
Southern Flying, 374 
Squirrel, Tamiasciurus hudsonicus, Short-tailed Shrew, Bla- 
rina brevicauda, Apparently Killed by Red, 456 
Squirrels, Sciurus carolinensis, in British Columbia, The Dis- 
tribution and Habitat Selection of Introduced Eastern 
Grey, 343 
St. John’s-wort, 14 
Starling, European, 71 
Starwort, Umbellate, 429 
Staurothele, 76 
drummondii, 78 
fissa, 78 
Steatoda bipunctata, 269 
Stellaria calycantha, 277 
longipes, 277 
umbellata, 418 
Stemonyphantes blauveltae, 265 
Stenamma diecki, 39 
impar, 39 
Steniolia, 6 
Stercorarius parasiticus, 211 
Stevens, S.D. and T.I. Wellicome. A Survey for Federally List- 
ed Grassland Birds at First Nations Reserves, 490 
Stevens, S.S., 459 
Stickleback, Brook, 221 
Threespine, 28 


INDEX TO VOLUME 119 


643 


Stictiella callista, 13 
corniculata, 13 
emarginata, 6 
evansi, 13 
formosa, \3 
pulchella, 13 
pulchella pulchella, 13 
pulchella serrata, \3 
serrata, 13 

Stictiella emarginata (Hymenoptera: Sphecidae)*, Nesting 
Behavior, Ecology, Seasonal and Geographic Distri- 
bution of the Sand Wasp, 6 

Stipa comata, 220 

Stocek, R'B, 351 

Stoneroller, Central, 18 

Stonewort, 16 

Strix nebulosa, 69 
varia, 367,379 

Stuckenia spp., 546 
filiformis, 548 
pectinata, 28,546 

Sturnella magna, 71 

Sturnus vulgaris, 7\ 

Styloctetor stativus, 265 

Sucker, White, 18,221,295 

Sundew, Great, 430 

Sunfish, Green, 21 
Longear, 18 
Orangespotted, 18 

Swallow, Barn, 70 
Cliff, 70 
Tree. 70 

Swallow-wort, Pale, 525 

Swallow-wort, Vincetoxicum rossicum, following Above- 
ground Tissue Loss: Implications for the Timing of 
Mechanical Control, Response of Pale, 525 

Swertia perennis, 417 

Swertia, Alpine Bog, 433 

Sylvester, R.M., S.E. Freeling, and C.R. Berry, Jr. First Rec- 
ord of the Plains Minnow, Hybognathus placitus, in 
Canada, 219 

Symphoricarpos accidentalis, 220 

Symphyotrichum ciliatum, 418 
lanceolatum ssp. hesperium var. hesperium, 439 
laurentianum, 556 
yukonense, 418 

Symphyotrichum laurentianum, aux [les-de-la-Madeleine, 
Protocole de suivi des populations d’aster du Saint- 
Laurent, 556 

Symplocarpus foetidus, 129 

Syrphus, 250 


Tachycineta bicolor, 70 
Tachysphex pechumani, 10 
similis, 10 
tarsatus, 10 
Taenia pisiformis, 324 
Tamarack, 255,404,574 
Tamias striatus, 323,578 
Tamiasciurus douglasii, 344 
hudsonicus, 43,121,323,343,456,591 
Tamiasciurus hudsonicus, Short-tailed Shrew, Blarina bre- 
vicauda, Apparently Killed by Red Squirrel, 456 
Tapinocyba bicarinata, 265 
exigua, 264 


644 


flagellata, 265 
lindrothi, 265 
minuta, 265 
scopuliferum, 262 
simplex, 265 
Tapinoma sessile, 39 
Tapinopa bilineata, 265 
Taraxacum spp., 552 
lyratum, 418 
Tardigrada, 586 
Taxidea taxus, 442,584 
Taxus canadensis, 457 
Teal, Blue-winged, 69,379,547 
Green-winged, 69,547 
Tegenaria atrica, 256 
domestica, 256 
duellica, 256 
Tennesseellum formicum, 265 
Tern, Black, 69,78 
Téte-de-boule, 361 
Tetragnatha caudata, 255 
elongata, 268 
extensa, 269 
laboriosa, 269 
vermiformis, 255 
versicolor, 269 
Tetramorium caespitum, 39 
Tetrao parviventris, 120 
tetrix, 120 
Texas, Swift Fox, Vulpes velox, Den Located Next to a Rail- 
road Track in Northwestern, 584 
Thamnophis sp., 292 
sirtalis, 458 
Thanatus formicinus, 268 
Theberge, J.B., Review by, 308 
Thellungiella salsuginea, 429 
Thelypteris palustris, 388 
Theonoe stridula, 269 
Theridion aurantium, 269 
differens, 269 
glaucescens, 269 
montanum, 269 
murarium, 269 
petraeum, 270 
pictum, 270 
sexpunctatum, 269 
tectum, 269 
Theridiosoma gemmosum, 255 
radiosum, 270 
Theridula emertoni, 270 
sphaerula, 270 
Thistle, Elk, 435 
Leafy, 435 
Thomomys spp., 175 
bottae, 551 
talpoides, 175,551 
Thomomys talpoides, in Central Alberta Alfalfa Fields, Body 
Weights of Adult and Juvenile Northern Pocket 
Gophers, 551 
Thomomys talpoides, Long-tailed Weasel, Mustela frenata, 
Movements and Diggings in Alfalfa Fields Inhabited 
by Northern Pocket Gophers, 175 
Thomson, E.R. Papillate Watermeal, Wolffia brasiliensis, in 
Eastern Ontario: An Addition to the Flora of Canada, 
137 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Thomson, J.A., 214 
Thorn, T.D., R.B. Emery, D.W. Howerter, J.H. Devries, and 
B.L. Joynt. Use of Radio-Telemetry to Test for Inves- 
tigator Effects on Nesting Mallards, Anas platyrhyn- 
chos, 541 
Thrasher, Brown, 71 
Thrush, Hermit, 70 
Swainson’s, 70 
Wood, 70 
Thuidium abietinum, 80 
Thuja occidentalis, 386,404 
plicata, 160,344 
Thymoites oleatus, 270 
Thyreosthenius parasiticus, 265 
Tibellus maritimus, 268 
Tick, Winter, 330 
Tigellinus tricornis, 266 
Tilia, 96 
americana, 369 
americana Var. americana, 96 
Timmins, C.A., 16 
Timoney, K.P. and J. Marsh. Lichen Trimlines in the Peace- 
Athabasca Delta: Variations in Flora, Form, and Dis- 
turbance Regime, 76 
Tiso vagans, 265 
Tit, Great, 120 
Greyheaded, 120 
Long-tailed, 120 
Marsh, 120 
Willow, 120 
Titmouse, Tufted, 374 
Toad, Western, 291 
Tofieldea coccinea, 277 
glutinosa ssp. brevistyla, 428 
pusilla, 277 
Tokaryk, T., Reviews by, 305,306 
Tolypella glomerata, 26 
Torreyochloa pallida var. fernaldii, 394 
Tortula ruralis, 80 
Toxostoma rufum, 71 
Trachynella nudipalpis, 266 
Tramea lacerata, 234 
Tree, Cherry, 344 
Trematode, 324 
Triadenum virginicum, 388 
Triantha glutinosa, 418 
Trichophorum alpinum, 418 
Trichopterna mengei, 264 
Trichostrongylus axei, 324 
Trientalis borealis, 240 
Triglochin maritimum, 278 
Tringa flavipes, 69,379 
melanoleuca, 379 
Trisetum spicatum, 278 
Triteleia grandiflora var. howellii, 248 
Trochosa terricola, 267 
terricola pratensis, 267 
Troglodytes aedon, 374 
troglodytes, 69 
Tropaeolum sp., 438 
Trout, Brook, 351 
Tscherter, U., 214 
Tsuga canadensis, 38,129,332,386 
heterophylla, 160,197,344 
mertensiana, 291 


2006 


Tunagyna debilis, 265 
Turdus migratorius, 70 
Tursiops sp., 217 
Turtle, Blanding’s, 389 
Painted, 389 
Twinberry, Black, 434 
Twinflower, 289 
Tympanuchus cupido, 507,515 
cupido cupido, 516 
cupido pinnatus, 523 
cupido X phasianellus, 507,515 
phasianellus, 507 
phasianellus campestris, 507,515 
phasianellus phasianellus, 507,515 
Tympanuchus cupido X phasianellus, Hybridization on Mani- 
toulin Island, Ontario, “Prairie Grouse”, 507 
Tympanuchus cupido x phasianellus, of Manitoulin Island, 
Ontario, Plumage and Internal Morphology of the 
“Prairie Grouse”, 515 
Typha spp., 221,403 
angustifolia, 16 
latifolia, 16,72,389,418 
Typhocrestus pygmaeus, 265 


Uesugi, A., 569 

Ulmus, 96 
americana, 96,370 
rubra, 370 

Ulothrix, 83 

Ulva intestinalis, 83 

Umbilicaria, 78 
deusta, 78 
muehlenbergii, 78 

Umbra limi, 21,361 

Umbre de vase, 361 

Ursus americanus, 50,164,298,323,339,381,574 
arctos, 339,449,574 

Ursus americanus, Ecology on the Northeast Coast of 
Labrador, Black Bear, 164 

Ursus arctos, from Coastal British Columbia, Observations of 
Autumn Courtship and Breeding in Brown Bears, 449 

Usui, M., P.G. Kevan, and M. Obbard. Pollination and Breed- 
ing System of Lowbush Blueberries, Vaccinium angus- 
tifolium Ait. and V. myrtilloides Michx. (Ericacaeae), 
in the Boreal Forest, 48 

Utricularia cornuta, 388 
macrorhiza, 137 
minor, 418 


Vaccinium spp., 44,48 
angustifolium, 48,239 
myrtilloides, 48 
uliginosum, 412 
vitis-idaea, 210,412 
vitis-idaea var. minus, 278 
Vaccinium angustifolium Ait. and V. myrtilloides Michx. 
(Ericacaeae), in the Boreal Forest, Pollination and 
Breeding System of Lowbush Blueberries, 48 
Vaccinium myrtilloides Michx. (Ericacaeae), in the Boreal 
Forest, Pollination and Breeding System of Lowbush 
Blueberries, Vaccinium angustifolium Ait. and, 48 
Vachon, J., B.F. Lavallée, et F. Chapleau. Charactéristiques 
dune population introduit du Grand brochet, Esox 
lucius, dans le lac Ramsay, Pare de la Gatineau, 
Québec, et impact sur l’ichtyofaune, 359 


INDEX TO VOLUME 119 


645 


Valerian, Sitka, 434 
Valeriana sitchensis, 418 
Valerianella locusta, 248 
Vallisneria americana, 16,546 
Veery, 70 
Ventre citron, 360 
pourri, 361 
rouge du nord, 360 
Verbena hastata, 418 
Vermivora peregrina, 7| 
ruficapilla, 70 
Vermontia thoracica, 265 
Veronica americana, 418 
Verrucaria, 76 
Vetch, American, 432 
Common, 248 
Viburnum alnifolium, 129 
cassinoides, 129 
nudum var. cassinoides, 240 
Vicia americana, 418 
sativa, 248 
Vincetoxicum rossicum, 525 
Vincetoxicum rossicum, following Aboveground Tissue Loss: 
Implications for the Timing of Mechanical Control, 
Response of Pale Swallow-wort, 525 
Vine, Dog-strangling, 525 
Viola adunca, 418 
macloskeyi ssp. pallens, 389 
praemorsa ssp. praemorsa, 248 
Violet, Hook-spur, 432 
Yellow Montane, 248 
Viorne a feuilles d’aulne, 129 
cassinoide, 129 
Vireo gilvus, 70 
olivaceus, 71 
Philadelphicus, 71 
solitarius, 71 
Vireo, Blue-headed, 71 
Philadelphia, 71 
Red-eyed, 71 
Warbling, 70 
Vole, 122,395 
Heather, 414 
Long-tailed, 445 
Meadow, 43,323,414,446 
Northern Red-backed, 445 
Red-backed, 323 
Rock, 412 
Southern Red-backed, 43,414 
Taiga, 446 
Vole, Microtus chrotorrhinus, in Labrador, A Range Exten- 
sion for the Rock, 412 
Volucella bombylans, 249 
Vulpes macrotis mutica, 134 
velox, 134,584 
vulpes, 120,323 
Vulpes macrotis mutica, Aggressive Behaviour Exhibited by 
a San Joaquin Kit Fox, 134 
Vulpes velox, Den Located Next to a Railroad Track in North- 
western Texas, Swift Fox, 584 


Wabasso cacuminatus, 265 
quaestio, 265 

Walckenaeria arctica, 266 
atrotibialis, 266 


646 


auranticeps, 266 
castanea, 266 
clavipalpis, 266 
communis, 266 
cuspidata, 266 
digitata, 266 
directa, 266 
exigua, 266 
karpinskii, 266 
lepida, 266 
pallida, 266 
palustris, 266 
redneri, 266 
spiralis, 266 
tricornis, 266 
Walmus borealis, 256 
Walnut, Black, 348 
Wandeler, A., 453 
Warbler, Black-and-white, 70 
Black-throated Blue, 70 
Black-throated Green, 70 
Cape May, 71 
Chestnut-sided, 71 
Connecticut, 70 
Magnolia, 70 
Mourning, 70 
Nashville, 70 
Northern Parula, 69 
Palm, 69 
Tennessee, 71 
Wilson’s, 69 
Yellow, 70 
Yellow-rumped, 70 
Warner, B.G., 64 
Warren, R., 453 
Wasp, Sand, 6 
Wasp, Stictiella emarginata (Hymenoptera: Sphecidae)*, 
Nesting Behavior, Ecology, Seasonal and Geographic 
Distribution of the Sand, 6 
Water-bear, 586 
Water-bears from the Rocky Mountains: A First Look at 
Alberta’s Tardigrade Fauna, 586 
Water-horehound, Tuberous, 288 
Watermeal, 137 
Watermeal, Wolffia brasiliensis, in Eastern Ontario: An Addi- 
tion to the Flora of Canada, Papillate, 137 
Water-milfoil, Verticillate, 432 
Watershield, 137 
Waterthrush, Northern, 69 
Waxwing, Cedar, 70 
Way, J.G. and R.L. Proietto. Record Size Female Coyote, 
Canis latrans, 139 
Weasel, Long-tailed, 175 
Short-tailed, 175,323 
Weasel, Mustela frenata, Movements and Diggings in Alfalfa 
Fields Inhabited by Northern Pocket Gophers, Tho- 
momys talpoides, Long-tailed, 175 
Weir, R.D., 296 
Wellicome, T.1., 490 
Wendland, J.M., 591 
Wersal, R.M., B.R. McMillan, and J.D. Madsen. Food Habits 
of Dabbling Ducks During Fall Migration in a Prairie 
Pothole System, Heron Lake, Minnesota, 546 
Whale, Beluga, 214 
Blue, 214 
Fin, 214 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Humpback, 214 
Minke, 214 
Whales, Balaenoptera acutorostrata, in the Saguenay- 
St. Lawrence National Marine Park, Novel Surface 
Feeding Tactics of Minke, 214 
Wheatgrass, Slender, 78 
Whitefaces, 234 
Whitefish, Atlantic, 294 
Whitefish, Coregonus huntsmani, from the Wild, First Record 
of Age 0+ Atlantic, 294 
Whitlow-grass, Alaska, 429 
Few-seeded, 430 
Gray-leaved, 430 
Rocky Mountain, 430 
Starflowered, 430 
Wigeon, American, 69 
Widgeon-grass, 28 
Widgeon Grass, balls in Saskatchewan Lakes, Occurrence, 
Composition and Formation of Ruppia, 114 
Willow, 65,77,132,135,175,186,208,220,441,574,591 
Arctic, 282 
Barren-ground, 428 
Bebb’s, 96 
Blue-green, 282 
Blueberry, 283 
Net-veined, 283 
Richardson’s, 283 
Round-leaf, 283 
Tea-leaf, 283 
Willowherb, Hornemann’s, 432 
Swamp, 288 
Wilsonia pusilla, 69 
Winchester, N.N., 192 
Wolf, 135,139,192,197,330,38 1,446,573 
Eastern Canadian, 339 
Gray, 1,101,197,442 
Newfoundland, 323 
Timber, 101 
Wolf, Canus lupus, Den and Associated Human Activity in 
the Southwestern Yukon Territory, An Ancient, 135 
Wolf, Canis lupus, Faeces Revise Occurrence Records for 
Mammals of British Columbia’s Coastal Archipelage, 
Facts from Faeces: Prey Remains in, 192 
Wolf, Canis lupus, near Icy Bay, Alaska, Attempted Predation 
of a Child by a Gray, 197 
Wolffia arhiza, 137 
borealis, 137 
brasiliensis, 137 
columbiana, 137 
papulifera, 137 
punctata, 137 
Wolffia brasiliensis, in Eastern Ontario: An Addition to the 
Flora of Canada, Papillate Watermeal, 137 
Wolverine, 120 
Wolves, Canis lupus, in Relation to Ungulate Kill Sites in 
Westcentral Alberta, Travel Rates of, 573 
Wolves, Canis lupus, in the Wild, Longevity and Productivity 
of Three, 446 
Wood-rush, Piper’s, 427 
Spiked, 427 
Woodcock, American, 69 
Woodland Star, Small-flowered, 248 
Woodley, E., Review by, 309 
Woodpecker, American Three-toed, 367 
Black-backed, 367 
Downy, 71,369,379 


2006 


Hairy, 71,369,379 
Pileated, 70,367,379 
Red-bellied, 369 
Red-headed, 369 
Woodpecker Nest Tree Characteristics in Upper Midwestern 
Oak Forests, 367 
Woodsia ilvensis, 278 
Woodsia, Rusty, 278 
Wool-grass, 137 
Worm, Meningeal, 334 
Wormwood, Dragon, 438 
Wren, House, 374 
Marsh, 69 
Winter, 69 
Wubana drassoides, 266 
pacifica, 266 


Xanthoparmelia somloénsis, 78 

Xanthoria elegans, 78 

x Ceratinops annulipes, 261 

Xerostictia, 6 

Xysticus canadensis, 270 
discursans, 270 
elegans, 270 
emertoni, 270 
keyserlingi, 270 
labradorensis, 270 
luctuosus, 270 
obscurus, 270 
triguttatus, 270 


Yarrow, 248,438 
Pearl, 438 
Yellowhammer, 120 


INDEX TO VOLUME 119 


647 


Yellowlegs, Greater, 379 
Lesser, 69,379 

Yellowthroat, Common, 69 

Yucca sp., 584 

Yucca, 584 

Yukon, First Confirmation of Cougar, Puma concolor, in the, 
580 

Yukon Territory, An Ancient Wolf, Canus lupus, Den and 
Associated Human Activity in the Southwestern, 135 

Yukon Territory VII, New Records of Vascular Plants in the, 
417 


Zannichellia spp., 548 
palustris, 28 
Zapus sp., 578 
Zea mays, 548 
Zelotes fratris, 259 
subterraneus, 259 
sula, 259 
Zenaida macroura, 71,379 
Zielinski, R., 537 
Zimmerling, J.R. Detectability of Non-passerines Using 
“Pishing” in Eastern Ontario Woodlands, 377 
Zimmerling, T.N. The Influence of Thermal Protection on 
Winter Den Selection by Porcupines, Erethizon dor- 
satum, in Second-Growth. Conifer Forests, 159 
Zonotrichia leucophrys, 71 
Zonotrichia albicollis, 70 
Zornella cultrigera, 266 
Zostera marina, 28 
Zygadenus venenosus, 248 
Zygiella atrica, 257 
montana, 257 
nearctica, 257 


648 


Index to Book Reviews 


Botany 

Blouin, G. An Eclectic Guide to Trees East of the Rockies, 
497 

Douglas, G.W., G.B. Straley, D. Meidinger and J. Pojar. Illus- 
trated Flora of British Columbia Volumes 1-8, 329 

Evans, C.L. The War on Weeds in the Prairie West: An Envi- 
ronmental History, 670 

Kershaw, L. Ontario Wild Flowers, 330 

Kirk, P.M., P.F Cannon, J.C. David and J.A. Stalpers. Ains- 
worth and Bisby’s Dictionary of the Fungi, 328 

Lesica, P. A Flora of Glacier National Park, Montana, 159 

Mitchell, R.S. and L. Danaher. Northeastern Fern Identifier, 
669 

More, D. and J. White. The Illustrated Encyclopedia of Trees, 
496 

Riley, J.L. Flora of the Hudson Bay Lowland and its Post- 
glacial Origins, 669 

Schnell, D.E. Carnivorous Plants of the United States and 
Canada: Second Edition, 496 

Environment 

Bierregaard, R.O. Jr., C. Gascon, T.E. Lovejoy, and R. Mes- 
quita. Lessons from Amazonia: The Ecology and Con- 
servation of a Fragmented Forest, 672 

DeGraaf, R.M. Trees, Shrubs, and Vines for Attracting Birds, 
160 

Dixon, T.F. City Wilds: Essays and Stories about Urban 
Nature, 498 

Donovan, T.M. and C.W. Welden. Spreadsheet Exercises in 
Conservation Biology and Landscape Ecology, 506 

Donovan, T.M. and C.W. Welden. Spreadsheet Exercises in 
Ecology and Evolution, 506 

D’ Orso, M. Plundering Paradise, 335 

Ducey, J.E. Birds of the Untamed West: The History of Bird- 
life in Nebraska, 1750 to 1875, 336 

Franklin, S.E. Remote Sensing for Sustainable Forest Manage- 
ment, 330 

Frelich, L.E. Forest Dynamics and Disturbance Regimes: Stu- 
dies from Temperate Evergreen-Deciduous Forests, 502 

Higgs, E. Nature by Design, 505 

Johnsgard, P.A. Great Wildlife of the Great Plains, 504 

Jones, H.G., J.W. Pomeroy, D.A. Walker and R.W. Hoham. 
Snow Ecology: An Interdisciplinary Examination of 
Snow-Covered Ecosystems, 332 

Larson, D., U. Matthes, and P. Kelly. Cliff Ecology: Pattern 
and Process in Cliff Ecosystems, 672 

Morris, W.F. and D.F. Doak. Quantitative Conservation Bio- 
logy: Theory and Practice of Population Viability 
Analysis, 507 

Nadkarni, N.M. and N.T. Wheelwright. Monteverde: Ecology 
and Conservation of a Tropical Cloud Forest, 334 

Pullin, A.S. Conservation Biology, 499 

Schwartzman, D. Life, Temperature, and the Earth, 498 

Song, S.J. Ecological Basis for Stand Management: A Sum- 
mary and Synthesis of Ecological Responses to Wild- 
fire and Harvesting in Boreal Forests, 500 

Suzuki, D. and A. McConnell. The Sacred Balance: A Visual 
Celebration of Our Place in Nature, 503 

Thomashow, M. Bringing the Biosphere Home, Learning to 
Perceive Global Environmental Change, 159 

Watt, A. The Last Island: A Naturalist’s Sojourn on Triangle 
Island, 505 

Wiese, F. Seabirds and Atlantic Canada’s Ship-Source Oil 
Pollution, 332 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Miscellaneous 

Arnaudin, M.P. A Bird in the Bush: The Story of the Province 
of Quebec Society for the Protection of Birds 1917- 
2002, 508 

Baker, D.B. John Keast Lord: Materials for a Life, 338 

Berry, M.F. The Dinosaur Filmography, 341 

Browne, J. Charles Darwin, The Power of Place, 340 

Debus, A.A. and D.E. Debus. Paleoimagery, The Evolution of 
Dinosaurs in Art, 341 

Hodgson, B. Naturalists: A Journal, 342 

Hodgson, B. Women Travelers: A Journal, 342 

King, J. Farley: The Life of Farley Mowat, 509 

Lawrence, R.D. The North Runner, 674 

Nicklen, P. and H. Brody. Seasons of the Arctic, 674 

Stroud, P.T. The Emperor of Nature: Charles-Lucien Bona- 
parte and his World, 337 

Suzuki, D. When the Wild Comes Leaping Up: Personal En- 
counters with Nature, 339,675 


Zoology 

Barter, M.A., Shorebirds of the Yellow Sea: Importance, 
Threats and Conservation Status, 321 

Behnke, R.J. Trout and Salmon of North America, 318 

Brooks, R.J., D. Strickland, and R.J. Rutter. Reptiles and Am- 
phibians of Algonquin Provincial Park, 490 

Byers, J.A. Built for Speed: A Year in the Life of Pronghorn, 
661 

Catling, P.M., C.D. Jones and P. Pratt. Ontario Odonata. Vol- 
ume 3 (including observations for the year 2001), 487 

Clements, J.F. Birds of the World: A Checklist, 322 

Collette, B.B. and G. Klein-MacPhee. Bigelow and Schroe- 
der’s Fishes of the Gulf of Maine, 319 

del Hoyo, J., A. Elliott, and J. Sargatal. Handbook of Birds of 
the World. Volume 8: Broadbills to Tapaculos, 663 

Downer, J. Weird Nature: An Astonishing Exploratioin of 
Nature’s Strangest Behavior, 158 

Eder, T. Mammals of Ontario, 487 

Ernst, C.H. and E.M. Ernst. Snakes of the United States and 
Canada, 495 

Fishpool, L.D.C. and M.I. Evans. Important Bird Areas in 
Africa and Associated Islands: Priority Sites for Con- 
servation, 664 

Gittleman, J.L., S.-M. Funk, D. MacDonald, and R.K. Wayne. 
Carnivore Conservation, 155 

Halliday, T. and K. Adler. Firefly Encyclopedia of Reptiles 
and Amphibians, 327 

Heintzelman, D. All-Weather Hawk Watcher’s Field Journal, 


326 

Holman, J.A. In Quest of Great Lakes Ice Age Vertebrates, 
494 

Houston, C.S. and W. Anaka. Birds of Yorkton — Duck Moun- 
tain, 668 


Jaramillo, A. Birds of Chile, 659 

Jones, K.A. Knowing Bass: The Scientific Approach to Catch- 
ing More Fish, 156 

Kays, R.W. and D.E. Wilson. Mammals of North America, 
488 

Knott, D.L. A Spring Expedition to the Falkland Islands and 
Antarctica, 320 

Lacey, E.A., J.L. Patton, and G.N. Cameron. Life Under- 
ground, the Biology of Subterranean Rodents, 157 

Leverton R. Enjoying Moths, 327 


2006 


Lieske, E. and R. Myers. Coral Reef Fishes: Indo-Pacific and 
Caribbean — Revised Edition, 662 

Love, M.S., M. Yoklavich and L. Thorsteinson. The Rock- 
fishes of the Northeast Pacific, 323 

Manly, B.J., L.L. McDonald, D.L. Thomas, T.L. McDonald 
and W.P. Erickson. Resource Selection by Animals, 
325 

Matthiessen, P. The Birds of Heaven: Travels with Cranes, 
158 

Maxwell, B.A., J.K. Werner, P. Hendricks, and D.L. Faith. 
Herpetology in Montana: A History, Status Summary, 
Checklists, Dichotomous Keys, Accounts of Native, 
Potentially Native, and Exotic Species, and Indexed 
Bibliography, 491 

Mecklenburg, C.W., T.A. Mecklenburg, and L.K. Thorstein- 
son. Fishes of Alaska, 151 

Riede, K. Global Register of Migratory Species (GROMS): 
Database, GIS Maps, and Threat Analysis, 154 


INDEX TO VOLUME 119 


649 


Rossi, J. and R. Rossi. Snakes of the United States and Can- 
ada: Natural History and Care in Captivity, 666 

Russell, C. and M. Enns. Grizzly Heart: Living without Fear 
among the Brown Bears of Kamchatka, 662 

Saffron, I. Caviar: The Strange History and Uncertain Future 
of the World’s Most Coveted Delicacy, 324 

Scott, J.M., S. Conant and C. Van Riper III. Evolution, Ecol- 
ogy, Conservation, and Management of Hawaiian 
Birds: A Vanishing Avifauna, 153 

Shirihai, H. The Complete Guide to Antarctic Wildlife, 489 

Sibley, D.A. Sibley’s Birding Basics, 151 

Szczerbak, N.N. Guide to the Reptiles of the Eastern Pale- 
arctic, 492 

Taylor, P. The Birds of Manitoba, 660 

Urquhart, E. and A. Bowley. Stonechats: A Guide to the 
Genus Saxicola, 667 

Zug, G.R., L.J. Vitt, and J.P. Caldwell. Herpetology: An 
Introductory Biology of Amphibians and Reptiles, 
Second Edition, 493 


650 


THE CANADIAN FIELD-NATURALIST 


Vol. 119 


Advice for Contributors to The Canadian Field-Naturalist 


Content 

The Canadian Field-Naturalist is a medium for the publi- 
cation of scientific papers by amateur and professional natu- 
ralists or field-biologists reporting observations and results 
of investigations in any field of natural history provided that 
they are original, significant, and relevant to Canada. All read- 
ers and other potential contributors are invited to submit for 
consideration their manuscripts meeting these criteria. The 
journal also publishes natural history news and comment items 
if judged by the Editor to be of interest to readers and sub- 
scribers, and book reviews. Please correspond with the Book 
Review Editor concerning suitability of manuscripts for this 
section. For further information consult: A Publication Policy 
for the Ottawa Field-Naturalists’ Club, 1983. The Canadian 
Field-Naturalist 97(2): 231-234. Potential contributors who 
are neither members of The Ottawa Field-Naturalists’ Club 
nor subscribers to The Canadian Field-Naturalist are encour- 
aged to support the journal by becoming either members or 
subscribers. 


Manuscripts 

Please submit, to the Editor, in either English or French, 
three complete manuscripts written in the journal style. 
The research reported should be original. It is recommended 
that authors ask qualified persons to appraise the paper before 
it is submitted. All authors should have read and approved it. 
Institutional or contract approval for the publication of the data 
must have been obtained by the authors. Also authors are ex- 
pected to have complied with all pertinent legislation regard- 
ing the study, disturbance, or collection of animals, plants or 
minerals. The place where voucher specimens have been de- 
posited, and their catalogue numbers, should be given. Lati- 
tude and longitude should be included for all individual local- 
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TABLE OF CONTENTS (concluded) Volume 119 Number 4 


Notes 
Caching behavior by wintering Northern Saw-whet Owls, Aegolius acadicus 
ARNOLD DEVINE and DWIGHT G. SNITH 


tirst confirmation of Cougar, Puma concolor, in the Yukon THOMAS S. JUNG and PHILIP J. MERCHANT 


sirst occurrence of the Round Goby, Neogobius melanostomus, in the St . Lawrence River 
at Cornwall, Ontario M. BRIAN MICKEY and ADRIENNE R. FOWLIE 


pt Fox, Vulpes velox, den located next to a railroad track in northwestern Texas 
KERRY L. NICHOLSON, BRADY K. MCGEE, and WARREN BALLARD 


ater-bears from the Rocky Mountains: A first look at Alberta's tardigrade fauna 
MATTHEW J. BOECKNER and HEATHER C . PROCTOR 


ortality of Little Brown Bats. Myotis lucifugus, in a rodent trap in the boreal forest 
THOMAS S . JUNG and BRIAN G . SLOUGH 


eavers, Castor canadensis, feeding on salmon carcasses: opportunistic use of a seasonally 
superabundant food source JEFFREY S. GLEASON, RYAN A. HOFFMAN, and JAMES M. WENDLAND 


ook Reviews | 

OOLOGY: Arrivals and Rivals: A Birding Oddity — Band-tailed Pigeon: Wilderness Bird at Risk — 
Monitoring Bird Populations Using Mist Nets: Studies in Avian Biology #29 — Portraits of the Bison: 
An Illustrated Guide to Bison Society — The Buffalo Wolf, Predators, Prey, and the Politics of Nature 
— Grassland Grouse and Their Conservation — Crows: Encounters with the Wise Guys — 
Antipredator Defenses in Birds and Mammals — For the Love of Insects — Rare Bird: Pursuing the 
Mystery of the Marbled Murrelet 


OTANY: Canada’ s Forests: A History 
NVIRONMENT: The Natural History of Bermuda — Saskatchewan Uncommon Views 


ISCELLANEOUS: Manly Hardy (1832-1910): The Life and Writing of a Maine Fur-buyer, Hunter, and 
Naturalist — Audubon in Edinburgh and his Scottish Associates — A History of Devonshire 
Ornithology — Mapper of Mountains: M. P. Bridgland in the Canadian Rockies, 1902-1930 


EW TITLES 


ews and Comment 
arine Turtle Newsletter (110) — Saving the Wild: RENEW 2005 — Canadian Species at Risk August 2005 


} e Ottawa Field-Naturalists' Club Awards for 2004 


dex to Volume 119 Compiled by LESLIE Copy 


idvice for Contributors to The Canadian Field-Naturalist 


jailing date of the previous issue 119(3): 30 November 2006 


2005 


578 
580 


582 


584 


586 


589 


591 


594 
603 
604 


606 
610 


613 


614 


618 


650 


THE CANADIAN FIELD-NATURALIST Volume 119 Number 4 


Articles 


Breeding and non-breeding range of Canada, Branta canadensis, and Cackling, 
Branta hutchinsii, Geese in the eastern Canadian Arctic 
MARK L. MALLORY, ALAIN J. FONTAINE, and HUGH Boyp 


A survey for federally listed grassland birds at First Nations Reserves 
ScoTT D . STEVENS, and Troy I . WELLICOME 


Addition to the flora of Canada? A specimen from the Arctic Archipelago, Northwest 
Territories, links two allopatric species of alkali grass, Puccinellia 
L. L. CONSAUL, L. J. GILLESPIE, and K. L. MACINNES 


“Prairie Grouse”, 7ympanuchus cupido X phasianellus, hybridization on 
Manitoulin Island, Ontario H. G. LUMSDEN 


Plumage and internal morphology of the “Prairie Grouse”, Tympanuchus cupido 
x phasianellus on Manitoulin Island, Ontario H. G. LUMSDEN 


Response of Pale Swallow-wort, Vincetoxicum rossicum, following aboveground tissue loss: 
implications for the timing of mechanical control 
Curtis J. MCKAGUE and NAOMI CAPPUCCINO 


Differential parental care by adult Mountain Plovers, Charadrius montanus 
STEPHEN J. DINSMORE and FRITZ L. KNOFT 


Body size distribution and frequency of anthropogenic injuries of Bluntnose Sixgill Sharks, 
Hexanchus griseus, at Flora Inlets, British Columbia 
ROBERT DUNBRACK and ROBERT ZIELINSKI 


Use of radio-telemetry to test for investigator effects on nesting Mallards, Anas platyrhynchos 
TERRI D. THORN, ROBERT B. EMERY, DAVID W. HOWERTER, 
JAMES H . DEVRIES, and BRIAN L. JOYNT 


Food habitats of dabbling ducks during fall migration in a prairie pothole system, 
Heron Lake, Minnesota RYAN M. WERSAL, BROCK R. MCMILLAN, and JOHN D. MADSEN 


Body weights of adult and juvenile Northern Pocket Gophers, Thomomys talpoides, 
in central Alberta alfalfa fields GILBERT PROULX 


Protocole de suivi des populations d' aster du Saint-Laurent, Symphyotrichum laurentianum, 
aux Iles-de-la-Madeleine GUILLAUME DE LAFONTAINE 


Tree recruitment limitation by introduced Snowshoe Hares, Lepus americanus, on Kent Island, 
New Brunswick TEVOR S . PETERSON, AKANE UESUGI, and JOHN LIGHTER 


Travel rates of Wolves, Canis lupus, in relation to ungulate kill sites in westcentral Alberta 
GERALD W. KUZYK, CHRISTOPH ROHNER, and FIONA K. A. SCHMIEGELOW 


(continued on inside back co 


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